Hydrobiologia 345: 149–184, 1997.

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c 1997 Kluwer Academic Publishers. Printed in Belgium.

Aquatic nematodes from Ethiopia VIII

Enoplids, with descriptions of Brevitobrilus fesehai n.sp. and B. tsalolikhini n.sp. (Enoplida:
Nematoda)

Eyualem Abebe1;2 & A. Coomans1

1
Instituut voor Dierkunde, Universiteit Gent, Ledeganckstraat 35, B-9000, Gent, Belgium
2
Bahir Dar Teachers’ College, Personal P.O. Box 223, Bahir Dar, Ethiopia

Received 21 November 1996; in revised form 11 February 1997; accepted 13 February 1997

Key words: Taxonomy, aquatic nematodes, Brevitobrilus, Epitobrilus, Tripyla, Ironus, new species, Ethiopia

Abstract

Two new and one known species of the genus Brevitobrilus, Epitobrilus setosus, Tripyla glomerans, and Ironus
ignavus and I. sphincterus are described from sediments of Lakes Shala, Tana and Ziway, and River Abbay,
Ethiopia. Brevitobrilus fesehai n. sp. can be recognized by a combination of characteristics: in having a narrow
anterior end, short cephalic and outer labial setae, long and bow-shaped spicules, the posteriormost supplement at a
distance of less than a spicule length from cloacal opening, numerous micropapillae, and terminal and subterminal
setae. B. tsalolikhini n.sp. can also easily be differentiated from all known species in the genus in having a
combination of characteristics: long cephalic and outer labial setae, short spicules without median stiffening piece,
the posteriormost supplement situated very close to the cloacal opening, supplements with straight duct that merges
with the dorsal wall of the ampulla, and elongate spermatozoa. The remaining five species are described in detail.
Also scanning electron microscopic pictures of B. fesehai n.sp., B. tsalolikhini n.sp., Epitobrilus setosus and Tripyla
glomerans are presented.

Introduction moment, excluding the two new species we describe

The genera Brevitobrilus and Epitobrilus were two
among the 13 genera proposed by Tsalolikhin (1981)
to accommodate the 72 species of the genus Tobrilus
Andrássy, 1959 sensu lato. He transferred 8 and 4
species to the former and latter genus, respective-
ly, from the genus Tobrilus Andrássy, 1959 sensu
lato. Tsalolikhin (1983) gave brief descriptions of the
species included in these genera.
Tsalolikhin (1992) proposed a subdivision of the
genus Brevitobrilus into two based on male supplement
characteristics: those species with their last supplement
close to the cloacal opening on the one hand and species
with their last supplement far (>1 spicule L) from the
cloacal opening on the other. Recent accounts of devel-
opments with regard to new and transferred species
in the genus are given in Ocaña et al. (1996). These
authors also described a new species from Spain. At the
here, the genus consists of 12 species of which three
are either originally described from or are known to
occur in Africa. An ubiquitous member of this genus,
B. graciloides, has an exceptionally well document-
ed distribution in most lakes of the southern part of
Ethiopia (Tsalolikhin, 1992).
The number of species in the genus Epitobrilus
has not changed since the genus was erected, but
the composition has. Tsalolikhin (1991) regarded
E. parvipapillatus (Kreis, 1932) Tsalolikhin, 1981
as species incertae sedis and transferred Eutobrilus
medius (Schneider, 1916) Tsalolikhin, 1981 to the
genus Epitobrilus based on stoma structure. Although
its individual species have a restricted distribution, Epi-
tobrilus is cosmopolitan as a genus.
Brzeski & Winiszewska-Slipiniska (1993) reviewed
the genera of the family Tripylidae. A large proportion
of the species in the family belong to the ubiquitous

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Article: hydr 3817 GSB: 7011158 Pips nr 136023 BIO2KAP
hydr3817.tex; 12/05/1997; 13:18; v.7; p.1
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genus, Tripyla Bastian, 1865. In addition these authors
listed 17 species in the genus of which three were new,
they also gave a key to the species of the genus and
redescribed 6 already known species.
Andrássy (1968) reviewed the genus Ironus Bastian
1865 and recognized only 10 of the by-then described
species to be valid, synonimizing the rest. Later, Ebsary
(1985) provided a key to the 20 species of the genus, i.e.
including those considered valid by Andrássy earlier as
well as to those described later. No new species of this
genus has been described in the last decade, therefore
Ebsary’s key could be considered still up-to-date and
is a useful tool in the identification of the species in
this genus. The most widely reported species in this
cosmopolitan genus is I. ignavus Bastian 1865.
In this paper, 3 species of the genus Brevitobrilus
of which 2 are new to science, Epitobrilus setosus,
Tripyla glomerans, and 2 species of the genus Ironus
are described in detail.
Description of the study area and material and
methods
We refer the reader to the first paper in the series
(Eyualem & Coomans, 1996) for detailed descriptions
of the study areas and accounts of the material and
methods employed in the study. All absolute measure-
ments in the tables are given in m.
Abbreviations
ABE = anterior body end
ABW = anal body width
Amph = amphid width
Amph W = amphidial fovea width
CBW = corresponding body width
CSL = cephalic setae length
Ex. pore = excretory pore
G1L = anterior genital branch
G2L = posterior genital branch
L = length
LM = light microscope
LRW = lip region width
LSL = labial setae length
MBW = maximum body width
n = number of specimens
NR = nerve ring from the anterior end
OLSPL = outer labial setiform-papillae length
PBE = posterior body end
Ph L = pharyngeal length (neck length)
RL = rectal length
SEM = scanning electron microscope
W = width
Descriptions of the species
1. Brevitobrilus graciloides (von Daday, 1908)
Tsalolikhin, 1981
Measurements
Tables 1 & 2 (Figures 1, 2 & 3)
Description
Population from Zegie
Female:
Body after fixation usually either curved ventrad
or undulated, sometimes straight or curved dorsad,
tapering towards the posterior end; maximum body
width anterior, at or slightly posterior to vulva. Cuticle
1.5–2.0 m thick, smooth under LM, finely striated
under SEM. Somatic setae numerous; distributed all
over the body. Below the cuticle is found a thin layer
of fine, dark-brown refractive particles. Their distrib-
ution uneven, with their highest concentration on the
ventral side, 25.0–40.0 m anterior and posterior to
vulva. Nine to 13 dorsolateral, type-II loxometanemes
on each side of the body; the first situated close to the
level of the nerve ring and the last in the tail; length of
frontal and caudal filament about 60 and 50 m respec-
tively. Metanemes overlapping in most part of the body,
overlap involving only two consecutive metanemes,
the caudal filament of the anterior metaneme and the
frontal filament of following metaneme. Crystalloid
bodies all over the body, situated most probably in the
pseudocoelom.
Lip region bluntly rounded, continuous with the
rest of the body. Sensilla prominent, in two circles:
an inner ring of six labial papillae and an outer ring of
four shorter (4.0–5.5 m) cephalic and six longer (7.0–
12.0 m) labial setae. Amphids stirrup-shaped, 14.1–
20.0% CBW; situated 0.5–0.7 LRW from ABE; fusus
amphidialis just behind fovea, as long as or slightly
longer than foveal width. Cheilostome distinct, with
sclerotized lining. Rest of stoma consisting of three
parts, an anterior and larger cup-shaped part followed
by two smaller ventrosublateral pouches, the first of the
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 Table 1. Measurements of females of Brevitobrilus graciloides (von Daday, 1908) Tsalolikhin, 1983 from three sites in Lake Tana viz. Gedero, Gelda and Zegie, from Lake Ziway, and
River Abbay.

L. Tana populations L. Ziway Abbay-1

Gedero (n = 10) Gelda (n = 10) Zegie (n = 10) (n = 10) (n = 2)

L 1529.0  96.8 (1413.0–1660.0) 1551.9 88.3 (1426.0–1748.0)

 1686.9  159.5 (1483.0–1990.0) 1622.9  99.7 (1535.0–1860.0) 1370, 1630
LRW 27.8  1.1 (26.0–29.0) 27.7 1.5 (24.0–29.0)
 27.8  1.5 (26.0–30.0) 25.8  2.2 (22.0–30.0) 20.0, 26.0
Amph–ABE 14.8  0.4 (14.0–15.0) 17.7 2.5 (14.0–22.4)
 16.9  2.1 (14.0–22.0) 16.3  0.7 (15.0–18.0) 14.0, 16.0
Amph W 4.5  0.5 (3.5–5.0) 4.3 0.3 (4.0–5.0)
 5.1  0.7 (4.0–6.5) 4.7  0.2 (4.0–5.0) 5.0, 6.0
CBW 29.8  1.5 (27.0–31.0) 30.5 0.8 (30.0–32.0)
 31.5  2.1 (28.0–35.0) 34.3  0.7 (33.0–35.0) 24.0, 28.0
Stoma L 15.8  0.4 (15.0–16.0) 14.9 0.9 (13.0–16.0)
 14.7  0.5 (14.0–15.5) 12.1  0.8 (11.0–13.0) 11.5, 15.0
Stoma W 12.8  0.2 (12.5–13.0) 11.2 0.8 (10.0–12.0)
 12.5  1.2 (10.0–15.0) 11.6  1.2 (10.0–13.0) 9.0, 11.0
Outer LSL 7.8  0.7 (7.0–9.0) 8.4 0.8 (7.0–10.0)
 8.3  1.4 (7.0–12.0) 6.8  0.4 (5.0–7.0) 5.0, 8.0
CSL 4.8  0.4 (4.0–5.0) 5.1 0.8 (4.0–7.0)
 4.9  0.4 (4.0–5.5) 4.3  0.4 (4.0–5.0) 3.0, 5.0
First tooth–ABE 18.0  0.6 (17.0–19.0) 18.5 1.9 (14.0–20.0)
 19.4  1.5 (17.0–22.0) 14.1  0.8 (13.0–15.0) 14.0, 18.0
First–second tooth 9.3  0.7 (8.0–10.0) 9.9 0.5 (9.0–11.0)
 10.3  0.8 (8.5–11.5) 9.4  0.9 (8.0–10.0) 9.0, 10.0
NR 105.8  5.0 (98.0–111.0) 112.1 4.9 (107.0–121.0)
 114.1  5.6 (105.0–123.0) 98.3  7.6 (89.0–115.0) 95.0, 118.0
Ph L 293.0  13.9 (270.0–310.0) 302.4 22.1 (263.0–342.0)
 303.4  24.0 (276.0–350.0) 308.6  33.3 (275.0–385.0) 234.0, 314.0
MBW 68.5  4.8 (63.0–77.0) 64.7 5.0 (61.0–76.0)
 65.2  4.2 (55.0–70.0) 50.1  2.6 (46.0–54.0) 63.0, 64.0
ABW 30.2  4.5 (24.0–37.0) 32.4 3.0 (26.0–37.0)
 32.7  7.1 (25.0–50.0) 25.3  1.2 (24.0–27.0) 29.0, 30.0
RL 39.8  3.2 (35.0–43.0) 35 2.5 (33.0–40.0)
 39.6  2.7 (35.0–45.0) 28.6  1.9 (25.0–31.0) 31.0, 38.0
G1L 252.5  35.2 (206.0–315.0) 289.0 32.0 (235.0–333.0)
 279.9  37.7 (230.0–353.0) 292.7  22.0 (254.0–314.0) 209.0, 318.0
G2L 289.2  45.6 (225.0–350.0) 287.7 30.8 (275.0–343.0)
 317.9  35.2 (274.0–370.0) 340.3  20.3 (314.0–364.0) 230.0, 320.0
V 684.5  71.5 (584.0–782.0) 715.3 43.7 (644.0–818.0)
 730.3  72.9 (629.0–874.0) 734.6  39.1 (688.0–809.0) 547.0, 790.0
Tail L 174.5  9.7 (159.0–188.0) 183.3 12.8 (164.0–207.0)
 208.0  23.3 (162.0–243.0) 160.3  8.9 (147.0–170.0) 161.0, 196.0
Egg L 59.6  5.7 (50.0–68.0) 45.0  11.1 (33.0–63.0) 41.0
Egg W 33.2  3.5 (27.0–37.0) 31.2  7.3 (21.0–43.0) 16.0

a 22.4  1.5 (19.3–23.7) 24.1  2.3 (20.5–28.7) 29.5  2.3 (23.5–30.2) 32.5  2.5 (29.9–38.0) 21.7, 25.5
b 5.2  0.4 (4.6–5.6) 5.1  0.2 (4.8–5.4) 5.6  0.2 (5.2–5.8) 5.3  0.4 (4.8–6.0) 5.2, 5.9
c 8.8  0.5 (8.2–9.7) 8.5  0.6 (7.9–9.6) 8.2  0.9 (7.0–10.4) 9.9  0.5 (9.2–10.6) 8.3, 8.5
c0 5.8  1.1 (4.6–7.8) 5.7  0.8 (5.0–8.0) 6.5  0.9 (4.9–8.1) 6.3  0.5 (5.5–7.0) 5.6, 6.5
V% 44.7  2.7 (40.4–47.5) 46.1  1.4 (43.8–48.1) 43.3  2.2 (38.2–46.0) 45.3  0.9 (43.5–46.7) 41.9, 48.8
NR% 36.2  2.9 (32.2–40.7) 37.1  2.0 (34.2–40.7) 37.7  1.8 (34.3–39.9) 31.9  1.3 (29.9–33.9) 37.5, 40.6
Amph–ABE/LRW 0.5  0.0 (0.5–0.6) 0.6  0.2 (0.5–0.7) 0.6  0.1 (0.5–0.7) 0.6  0.2 (0.6–0.7) 0.5, 0.8
Amph W% CBW 14.9  1.5 (13.1–18.8) 13.9  0.9 (13.3–15.6) 16.2  1.7 (14.1–20.0) 13.6  0.9 (12.1–14.3) 17.8, 25.0

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Table 2. Measurements of males of Brevitobrilus graciloides (von Daday, 1908) Tsalolikhin, 1983 from Gedero and Zegie, Lake Tana, from Lake Ziway and River Abbay.

L. Tana populations
Gelda (L. Tana) (n = 10) Zegie (L. Tana) (n = 10) Lake Ziway (n = 10) Abbay-1 (n = 3)

L 1357.7  77.3 (1209.3–1432.0) 1514.4  186.1 (1288.0–1882.0) 1502.6  19.35 (1349.0–1628.0) 1246.0, 1350.0, 1452.0
LRW 22.2  1.2 (20.0–24.0) 23.2  1.9 (21.0–27.5) 21.1  1.8 (19.0–24.0) 20.0, 22.0, 23.0
Amph–ABE 13.0  0.0 (13.0–13.0) 14.2  1.7 (11.0–16.0) 14.8  1.2 (14.0–15.0) 13.0, 15.0, 15.0
Amph W 4.8  0.2 (4.0–5.0) 5.1  0.7 (4.0–6.0) 5.0  0.0 (5.0–5.0) 5.0, 5.0, 7.0
CBW 23.3  1.8 (22.0–25.0) 26.1  1.6 (23.0–29.0) 27.0  1.1 (26.0–28.0) 23.0, 24.0, 25.0
Stoma L 13.3  0.7 (12.0–14.0) 12.4  0.9 (10.5–14.0) 11.0  0.5 (10.0–12.0) 12.0, 13.0, 13.0
Stoma W 9.2  0.7 (8.0–10.0) 10.3  0.7 (9.5–11.5) 8.8  0.7 (8.0–10.0) 8.0, 9.0, 9.0
Outer LSL 7.2  0.7 (6.0–8.0) 7.3  1.4 (6.0–10.0) 6.5  1.4 (6.0–7.0) 6.0, 6.0, 7.0
CSL 5.7  0.5 (5.0–6.0) 4.9  0.5 (4.0–6.0) 4.5  0.4 (4.0–5.0) 4.0, 4.0, 5.0
First tooth–ABE 16.2  0.7 (15.0–17.0) 17.0  0.7 (16.0–18.0) 13.8  1.5 (12.0–15.0) 15.0, 15.0, 16.0
First–second tooth 9.0  0.5 (8.0–10.0) 10.3  0.9 (9.0–12.0) 8.1  0.1 (8.0–9.0) 9.0, 10.0, 10.0
NR 101.0  1.5 (98.0–102.0) 109.2  6.3 (102.0–122.0) 91.8  6.7 (81.0–106.0) 87.0, 96.0, 96.0
Ph L 267.8  4.3 (261.0–274.0) 281.5  17.4 (250.0–312.0) 289.7  18.5 (265.0–326.0) 208.0, 237.0, 258.0
MBW 50.7  2.4 (48.0–54.0) 54.5  3.4 (49.0–62.0) 39.6  2.1 (37.0–44.0) 50.0, 52.0, 53.0
ABW 29.3  1.6 (27.0–32.0) 32.4  3.7 (27.0–40.0) 25.9  1.7 (23.0–28.0) 30.0, 31.0, 35.0
Spicule L 36.8  2.1 (34.0–41.0) 39.8  3.3 (35.0–46.0) 32.0  2.5 (27.0–35.0) 34.0, 39.0, 40.0
Gubernaculum L 14.7  2.6 (10.0–17.0) 16.4  3.7 (12.0–22.0) 15.8  1.9 (13.0–19.0) 13.0, 16.0, 17.0
Distance between supplements
First–second 37.1 2.1 (35.0–40.0)
 34.6  3.4 (27.0–38.0) 34.0  1.6 (33.0–35.0) 30.0, 35.0, 36.0
Second third 24.3 1.0 (23.0–25.0)
 24.4  2.0 (22.0–27.0) 16.2  1.3 (15.0–29.0) 17.0, 25.0, 26.0
Third–fourth 35.0 2.1 (33.0–38.0)
 34.9  6.0 (25.0–45.0) 30.2  1.1 (26.0–35.0) 22.0, 28.0, 41.0
Fourth–fifth 34.6 3.0 (30.0–40.0)
 33.5  5.1 (23.0–40.0) 36.5  2.1 (28.0–44.0) 25.0, 33.0, 35.0
Fifth–sixth 43.5 4.7 (37.0–54.0)
 44.9  8.7 (33.0–60.0) 48.5  1.7 (38.0–50.0) 30.0, 37.0, 45.0
Sixth–cloaca 37.5 1.9 (34.0–40.0)
 38.6  6.1 (29.0–48.0) 32.1  4.7 (22.0–37.0) 28.0, 30.0, 30.0
Tail L 132.7 7.0 (123.0–146.0)
 145.7  9.6 (132.0–158.0) 115.0  11.1 (96.0–131.0) 102.0, 135.0, 140.0

a 27.2  1.6 (25.2–29.8) 27.8  3.1 (23.0–34.0) 38.1  3.2 (32.8–41.7) 23.5, 27.0, 27.9
b 5.1  0.3 (4.6–5.3) 5.4  0.5 (4.8–6.3) 5.2  0.2 (5.0–5.5) 5.6, 5.7, 6.0
c 10.4  0.6 (9.6–11.0) 10.4  1.0 (8.2–12.1) 13.2  1.1 (11.6–15.9) 10.0, 10.4, 12.2
c0 4.5  0.2 (4.4–4.8) 4.6  0.6 (3.9–5.7) 4.4  0.3 (4.0–4.9) 3.4, 4.0, 4.4
NR% 37.6  0.7 (36.6–38.3) 38.9  2.1 (36.1–42.3) 31.7  1.0 (29.6–32.8) 37.2, 40.5, 41.8
Amph–ABE/LRW 0.6  0.0 (0.6–0.6) 0.6  0.1 (0.5–0.7) 0.7  0.0 (0.7–0.7) 0.6, 0.7, 0.8
Amph W% CBW 20.1  2.0 (16.0–22.1) 19.7  3.2 (13.8–23.9) 18.5  0.8 (17.9–19.2) 20.8, 21.7, 28.0

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Figure 1. B. graciloides, female, A & D: Median and surface views respectively of the anterior end; B: Reproductive system; C: Anterior part
of pharynx showing positions of the nerve ring and excretory pore; E: Surface view of the pharyngo-intestinal junction showing coelomocytes
and pericardial gland cells; F & H: Median and surface views respectively of rectal area and tail; G: Entire female; I: Median view of
pharyngo-intestinal junction.

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154

Figure 2. B. graciloides, male, A–C: Tail and male reproductive system till the level of fine granulation; D & F: Median and surface views
respectively of the anterior end; E: Surface view of the pharyngo-intestinal junction showing coelomocytes and pericardial gland cells; G:
Pharyngeal region; H: Entire male; I: Testes; J: Surface view of tail.

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Figure 3. SEM pictures, B. graciloides, A–E female, A: Head-on view; B–D: Longitudinal sections of the anterior end showing, B: the
cup-shaped part of the stoma, the first, right, ventrosublateral pouch and its associated tooth (arrow); C: the cup-shaped part of the stoma, the
dorsal gland opening (arrow), the second, left ventrosublateral pouch and its associated tooth; D: the cup-shaped part of the stoma, the second,
left, ventrosublateral pouch and its associated tooth (arrow); E: Lateral view of anterior (arrowhead pointing at amphid opening); F–G, male, F:
Ventral view showing the last two ventromedian supplements and the cloacal opening; G: A ventromedian supplement. (Scale bar, A–F = 10 m;
G = 1 m).

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156
latter on the right the second on the left side (Figure 3B–
D). Pouch wall well sclerotized; each pouch is supplied
with a well sclerotized tooth. Cup-shaped part 1.0–
1.5 times longer than wide. First tooth situated 17.0–
22.0 m from ABE, the second tooth situated 8.5–
11.5 m from the first.
Pharynx cylindrical and muscular. Dorsal gland
opening at the level posterior to the first ventrosublat-
eral pouch (Figure 3C). Nerve ring prominent, at about
one-third to two-fifths of pharyngeal length. Cardia
18–20  20–25 m. Three prominent (16–20  25–
28 m) cells are situated at the junction of the car-
dia with the pharynx base. On each side of the body,
close to the base of the pharynx are situated two highly
granular coelomocytes(?). One coelomocyte is situat-
ed anterior to the level of the cardia while the second
is posterior to the level of the base of pharynx; they
are longer than wide. Two to three relatively smaller
coelomocytes are situated dorsal to the rectum. Excre-
tory pore faintly detectable, situated 100.0–115.0 m
from the ABE. Ventral gland cells not observed. Intesti-
nal wall cells filled with brown granules; with a distinct
brush-border; posteriorly leading to 35.0–45.0 m long
rectum.
Reproductive system didelphic, amphidelphic.
Gonoducts situated ventral to the intestine; direction
of flexure variable. Spermatheca distinct. Uterus sup-
plied with well developed, externally transversely stri-
ated muscular wall. Ovejector with wide lumen; its
wall internally lined by highly plicate layer. Vagina
15.0–20.0% CBW; with inconspicuous sphincter mus-
cle. Vaginal wall without plication. Vulva anterior to
middle. Eggs longer than wide.
Tail elongate-conoid, about 6.0 rectal lengths long,
almost always straight, gradually narrowing for most
part with the last about 15 m long posterior end being
slightly swollen and club-shaped. Two to nine setae on
tail; terminal setae present in about 80% of the speci-
mens. Caudal glands three, prominent, arranged more
or less in tandem; terminating in a short, tubular spin-
neret outlet that protrudes out of the otherwise blunt tail
end. Spinneret outlet directed towards the ventral side
of the tail terminus, never exactly terminally directed.
Populations from Gedero, Gelda, L. Ziway & Abbay-1
In most respects these four populations are similar to
each other as well as to the population from Zegie.
However, the population from L. Ziway has a slightly
shorter stoma. Also one of the two specimens from
Abbay-1 was smaller than specimens from other sites.
Population from Zegie
Male:
Resembling female in many respects except the fol-
lowing. Body after fixation more undulated and coiled
than in female, posterior part strongly curved either
ventrad or dorsad; slimmer than female, maximum
body width often at the level where the two testis join
the gonoduct. Seven to 10 metanemes on each side of
the body. Only a single coelomocyte is present poste-
rior to the level of the cardia.
Reproductive system diorchic with the posterior
testis reflexed posteriad; situated ventral to the intes-
tine. Spicules slightly curved, each with a well scle-
rotized median stiffening piece. Part of gubernaculum
(corpus) enclosing dorsal side of both spicules, and its
curved cuneus passing in between the spicules; corpus
more or less straight. Six ejaculatory glands present on
each side; their ducts join to form a common duct one
on each side ventral to the anterior end of each spicule;
exact position where common duct opening is situated
is inconspicuous. Vas deferens long, extending anterior
to mid-body; component cells variable in appearance
from one part to the other: cells are bigger and hya-
line closer to cloaca, are partly filled with fine granules
slightly anterior to the level of the fourth supplement
till the level of the third supplement, are filled with
larger granules between the level of the second and
first supplement (= anterior most supplement), and are
more hyaline ventrally but filled with fine granules dor-
sally anterior to the level of the first supplement until
the well developed sphincter. The latter well demar-
cated, about 80 m long. The granules are fine in cells
situated in a region 150–170 m long, in front of the
sphincter but become extremely fine more anteriorly.
Spermatozoa more or less globular each with a dark,
usually centrally positioned spot.
Ventral supplements of two types: six prominent,
laterally bulb-shaped, 9.0–12.0 m long and 10.0–
14.0 m wide supplements and up to about 80 incon-
spicuous, laterally papilloid supplements [referred to
as micropapillae by Tsalolikhin (1993)] situated in
between the bulb-shaped supplements and anterior to
the first and posterior to the last supplement. Each
bulb-shaped supplement has 2–3 internal tubular struc-
tures (ducts) that are curved anteriad to fuse with the
anterior wall of the ampulla. Under SEM these sup-
plements appear to be composed of two concentric
rings with a plug-like structure (seta?) in the centre.
The plug-like structure may be protrusible, for we
have observed it to be of varying length in different
hydr3817.tex; 12/05/1997; 13:18; v.7; p.8

supplements. Distance between bulb-shaped supple-
ments variable, the shortest distance in most speci-
mens (22.0–27.0 m) was between the second and third
supplement while the longest distance (33.0–60.0 m)
in most specimens was between the fifth and sixth
supplement (Table 2). Micropapillae in some speci-
mens whose posterior part curved ventrad were two-
humped, this appearance however was not observed
or was rarely observed in specimens whose posterior
part was curved dorsad. Under SEM these micropapil-
lae appear to be ventral bands, about 12–15 m wide
and 1–2 m long, separated from each other by a line.
Number of micropapillae variable: 27–38 anterior to
the first supplement, 6–10 between the first and sec-
ond supplement, 5–12 between the second and third
supplement, 7–10 between the third and fourth supple-
ment, 6–10 between the fourth and fifth supplement,
8–16 between the fifth and sixth supplement, and 2–11
between the sixth supplement and cloacal opening.
Tail conoid and continuously narrowing till about
half of its length, more or less cylindrical in its posterior
half. Six to 8 setae on each side of tail, terminal seta
observed in about 90% of the specimens.
Population from Gelda, L. Ziway & Abbay-1
Except for the slimmer body and slightly more anteri-
orly situated nerve ring in the males from L. Ziway, the
males from the three sites are similar in every respect
to the population from Zegie.
Comparisons and rationale for identification
Tsalolikhin (1992) subdivided the genus Brevitobrilus
Tsalolikhin, 1981 into two groups: the first group
includes those species whose last supplement (sup-
plement closest to cloacal opening) is situated equal or
more than one spicule length from the cloacal open-
ing, while the second includes the ones that have
their last supplement situated less than one spicule
length from the cloacal opening. Since this distance
can easily be affected by the extent of coiling, it is
advisable to consider specimens with straight posteri-
or body. Our populations fall under the first group.
Comparison of our populations with those species
that fall under the first group showed that they come
close to B. graciloides (von Daday, 1908) Tsalolikhin,
1981 and B. stefanskii (Micoletzky, 1925) Tsalolikhin,
1981. Tsalolikhin (1992) designated a neotype from
L. Bishoftu, Ethiopia, for the former species and dis-
cussed the differences of the two species. He pointed
out that the two species can be differentiated from each
157
other by the length of the ejaculatory duct (that part of
the vas deferens posterior to the sphincter) compared to
the distance of the first bulb-shaped supplement from
the cloacal opening, by the number of micropapillae
anterior to the first supplement and the shape of these
micropapillae. It should be noted, however, that Tsalo-
likhin used the word ‘ductus ejaculatorius’ as a struc-
ture different from vas deferens as it could be under-
stood from his discussion on B. granatensis (Ocaña &
Zullini, 1988) Tsalolikhin, 1992 (Tsalolikhin, 1992).
Nevertheless, his mention of ‘... a different position of
the junction between vas deferens and ductus ejacu-
latorius and the lack of distinct sphincter between the
two ....’ indicates that he was referring to the part of the
vas deferens posterior to the sphincter. The sphincter
in males of our population was situated far anterior to
the first supplement and the number of micropapillae
was more than 27. Therefore our populations are closer
to B. graciloides, for these, according to Tsalolikhin
(1992), are two of the distinguishing characteristics
of the species. Tsalolikhin (1992) also characterised
B. graciloides by having terminal seta. But, although
most of our specimens of both sexes have terminal seta,
some specimens lack it. When seen as percentage, the
number could seem low, but is significant when seen
with respect to the number of specimens we studied
(>150). Whether this is because it is easily broken off
or due to its absence is not clear. Despite this singular
difference, our populations are similar to B. graciloides
and are identified as such.
Ocaña et al. (1996) transferred Tobrilus siculus Vin-
ciguerra & Zullini (1991) and T. sardus Vinciguerra &
Zullini (1991) to the genus Brevitobrilus. Our study
of type materials and original description of these two
species showed that their stoma structure and male
supplement morphology confirms the view of Ocaña
et al. (1996) in that they belong to the genus Brevi-
tobrilus and not to Tobrilus. The differences between
B. siculus and B. graciloides are pointed out in the orig-
inal description of the former (Vinciguerra & Zullini,
1991). Both transferred species fall into Tsalolikhin’s
species group‘B’.
B. graciloides was reported from Ethiopia first by
Filipjev (1931) under the name Trilobus graciloides.
He recorded it from six localities. Zullini (1988) and
Tudorancea & Zullini (1989) reported it from four rift
valley lakes viz. Lakes Abiyata, Awasa, Langano and
Ziway, and L. Koka under the name Tobrilus africanus.
The current study shows that this species is also found
far from the Ethiopian rift system.
hydr3817.tex; 12/05/1997; 13:18; v.7; p.9
158
Our study of specimens used by Zullini (1988)
showed that our populations are similar to his pop-
ulations.
2. Brevitobrilus fesehai n.sp.
Measurements
Table 3 (Figures 4, 5 & 6)
Type habitat and locality
Eight males and 4 females from surface to 3.0 cm of
sediment at 0.5 m depth, Gedero, L. Tana.
Additional collection
Four males and 1 female from surface to 3.0 cm sedi-
ment at 0.5 and 1.5 m depths, Zegie, L. Tana.
Type specimens
Holotype male (N 3917) with 3 female paratypes
deposited in the collection of the University of Ghent,
Instituut voor Dierkunde, Gent, Belgium. A male
paratype distributed to each of the following collec-
tions: USDA Nematode Collection, Beltsville, Mary-
land, USA; Nematode Collection, International Insti-
tute of Parasitology, St. Albans, UK; Nematode Col-
lection of Landbouwhogeschool, Wageningen, the
Netherlands; Australian Collection at Canberra.
Etymology
The species is named after Mr Feseha Abebe whose
help during sample collection was immense.
Description
Type population
Female:
Body after fixation straight (two specimens) or
undulated (two specimens), tapering towards both
ends; maximum body width anterior, at or slightly
posterior to vulva. Cuticle 2.0–3.0 m thick, its layers
conspicuous, a thinner, lighter outer and slightly thick-
er and darker inner layer; smooth under LM. Somatic
setae few. Below the cuticle is found a thin layer of
fine, dark-brown refractive particles. Six to nine dor-
solateral, type-II loxometanemes on each side of the
body; the first situated close to the level of the nerve
ring and the last in the tail; length of frontal and caudal
filament about 45 and 60 m, respectively. Metanemes
overlapping in most part of the body, overlap involving
only two consecutive metanemes, the caudal filament
of the anterior metaneme and the frontal filament of
following metaneme. Crystalloid bodies all over the
body, situated most probably in the pseudocoelom.
Lip region narrow, bluntly rounded, continuous
with the rest of the body. Sensilla in two circles: an
inner ring of six labial papillae and an outer ring of
four short (2.5–3.0 m) cephalic and six slightly longer
(3.0–4.0 m) labial setae. Amphids stirrup-shaped
with transverse oval aperture, 19.6–25.0% CBW; sit-
uated 0.8–1.0 LRW from ABE; fusus amphidialis just
behind fovea, as long as or slightly longer than fovea.
Cheilostome distinct, with weakly sclerotized lining.
Rest of stoma consisting of three parts, an anterior and
larger cup-shaped part followed by two smaller ventro-
sublateral pouches of which the first is situated on the
right and the second on the left side. Pouch wall well
sclerotized; each pouch is supplied with a well sclero-
tized tooth. Cup-shaped part 1.4–1.7 times longer than
wide. First tooth situated 12.5–16.0 m from ABE, the
second tooth situated 8.0 m from the first.
Pharynx cylindrical and muscular. Nerve ring
prominent, at about one-third to two-fifths of pharyn-
geal length. Cardia 13–15  16–18 m. Three promi-
nent (13–15  19–22 m) cells situated on the junction
of the cardia with the pharynx base. On the dorsal side,
posterior to these cells is situated one highly granular
coelomocyte (?); it is longer than wide. Excretory pore
situated about 85.0 m from the ABE. Ventral gland
cells not observed. Cells of intestinal wall filled with
brown granules. Intestine leading to 28.0–35.0 m long
rectum.
Reproductive system didelphic, amphidelphic.
Gonoducts situated ventral to the intestine; direction
of flexure variable. Spermatheca indistinct. Uterus sup-
plied with well developed, externally transversely stri-
ated muscular wall. Ovejector with wide lumen; its
wall internally lined by highly plicate layer. Vagina
about 10.0% CBW; with inconspicuous sphincter mus-
cle. Vaginal wall without plication. Vulva anterior to
or at mid-body. Up to 4 eggs in the uterus.
Tail elongate-conoid, about 5.0 rectal lengths long,
almost always straight, gradually narrowing for most
part with the last about 15 m long posterior end being
slightly swollen and club-shaped. Four to seven setae
on tail; terminal setae absent. Caudal glands three,
hydr3817.tex; 12/05/1997; 13:18; v.7; p.10
 159
Table 3. Measurements of males and females of Brevitobrilus fesehai n. sp. from Gedero and Zegie, Lake Tana.

Type population from Gedero Population from Zegie

paratype females Holotype paratype males female males
(n = 4) male (n = 8) (n = 1) (n = 4)

L 
1236.0 141.8 (1041.0–1432.0) 1090.0 1096.3  91.7 (988.0–1286.0) 844.0 1088.0  58.9 (997.0–1141.0)
LRW 
18.3 1.9 (15.0–20.0) 17.0 17.4  0.7 (16.0–18.0) 17.0 17.1  0.5 (16.5–18.0)
Amph–ABE 
15.9 0.9 (15.0–17.0) 13.0 14.0  0.9 (13.0–15.0) 19.0 14.5  0.6 (13.5–15.0)
Amph W 
5.1 0.9 (4.0–6.0) 7.0 6.1  0.4 (5.5–7.0) 4.5 5.8  0.6 (5.0–6.5)
CBW 
23.0 2.5 (19.0–26.0) 22.0 20.9  1.1 (19.0–22.0) 22.0 21.0  1.0 (20.0–22.0)
Stoma L 
12.8 0.4 (12.0–13.0) 13.0 11.9  0.7 (11.0–13.0) 12.0 10.0  0.7 (9.0–11.0)
Stoma W 
8.0 0.7 (7.0–9.0) 7.0 6.9  0.5 (6.0–7.5) 7.0 6.9  0.4 (6.5–7.5)
Outer LSL 
3.8 0.4 (3.0–4.0) 4.0 3.7  0.2 (3.5–4.0) 4.0 3.4  0.2 (3.0–3.5)
CSL 
2.9 0.2 (2.5–3.0) 3.0 3.0  0.0 (3.0–3.0) 3.0 3.0  0.0 (3.0–3.0)
First tooth–ABE 
14.6 1.3 (12.5–16.0) 14.0 13.3  0.5 (13.0–14.0) 17.0 13.4  0.6 (13.0–14.5)
First–second tooth 
8.0 0.0 (8.0–8.0) 8.0 8.1  0.3 (8.0–9.0) 9.0 8.0  0.4 (7.5–8.5)
NR 
91.5 5.4 (85.0–100.0) 90.0 88.3  3.7 (82.0–92.0) 102.0 92.3  3.3 (87.0–95.0)
Ph L 
233.8 22.9 (203.0–260.0) 210.0 216.1  13.0 (189–236) 218.0 215.5  15.0 (191.0–230.0)
MBW 
54.3 7.2 (44.0–62.0) 52.0 45.1  2.4 (42.0–48.0) 50.0 42.8  1.6 (40.5–45.0)
ABW 
29.8 2.9 (26.0–33.0) 33.0 28.9  2.9 (24.0–32.0) 26.0 30.5  2.1 (29.0–34.0)
RL 
31.5 2.7 (28.0–35.0) 30.0
Tail L 
156.3 8.2 (145.0–165.0) 116.0 118.6  7.5 (106–133) 140.0 113.8  8.2 (102.0–125.0)
G1L 
185.3 54.7 (100.0–245.0) 116.0
G2L 
185.8 63.1 (80.0–235.0) 148.0
V 
563.5 72.5 (445.0–626.0) 430.0
Spicule L 48.0 43.7  2.0 (41.0–46.0) 44.0  2.5 (41.0–48.0)
Gubernaculum L 17.0 16.9  2.7 (12.0–20.0) 15.5  2.7 (13.0–20.0)
Distance between supplements
First–second 35.0 31.1  4.7 (23.0–38.0) 26.9  3.2 (22.5–30.0)
Second–third 37.0 29.6  5.2 (22.0–38.0) 22.5  4.0 (17.5–27.5)
Third–fourth 34.0 28.6  4.0 (22.0–34.0) 25.5  5.8 (18.5–31.5)
Fourth–fifth 30.0 27.0  2.9 (23.0–31.0) 21.0  5.0 (15.0–26.0)
Fifth–sixth 48.0 41.6  9.3 (27.0–53.0) 32.8  6.9 (25.0–43.0)
Sixth–cloaca 31.0 29.6  3.2 (25.0–35.0) 26.6  4.9 (20.0–33.5)
a 23.1  3.2 (19.9–28.1) 21.0 24.3  1.8 (22.2–27.4) 16.9 25.5  1.8 (23.2–28.1)
b 5.3  0.4 (4.7–5.8) 5.2 5.1  0.2 (4.7–5.4) 3.9 5.1  0.2 (4.7–5.3)
c 7.9  0.7 (7.2–8.7) 9.4 9.3  1.1 (7.4–10.6) 6.0 9.6  0.4 (9.1–10.1)
c0 5.3  0.4 (4.8–5.8) 3.5 4.2  0.7 (3.3–5.3) 5.4 3.7  0.4 (3.4–4.3)
V% 45.6  3.7 (42.7–51.9) 50.9
NR% 39.4  2.8 (34.6–41.9) 47.6 40.9  2.2 (36.4–43.4) 46.8 
42.9 1.6 (41.3–45.5)
Amph–ABE/LRW 0.9  0.1 (0.8–1.0) 0.8 0.8  0.1 (0.7–0.9) 1.1 
8 0.1 (0.8–0.9)
Amph W% CBW 22.5  2.3 (19.6–25.0) 31.8 29.1  1.5 (27.3–31.8) 20.5 
27.3 1.6 (25.0–29.5)

prominent; terminating in a short, tubular spinneret amphids were situated slightly more posterior than in
outlet that opens perfectly terminal on the blunt tail the type population. Despite these differences the spec-
end. imen shows close similarity to the type population in
every other respect.
Specimen from Zegie
The single specimen from Zegie was shorter, and had Type population
lower values for the ratios ‘a’, ‘b’ and ‘c’, and the Male:

hydr3817.tex; 12/05/1997; 13:18; v.7; p.11

160

Figure 4. B. fesehai n. sp., female, A: Anterior part of pharyngeal region till nerve ring and excretory pore; B: Rectal region and tail; C: Entire
female; D: Reproductive system; E: Surface view of anterior end; F: Surface view of pharyngo-intestinal junction.

Resembling female in many respects except the
following. Body after fixation more undulated and
coiled than in female, posterior part strongly curved
either ventrad or dorsad; maximum body width often
at the level of the testis. Amphids wider than in
females, 27.3–31.8% CBW, fusus amphidialis short-
er than fovea. Excretory pore situated at about 90 m
from ABE.
Reproductive system diorchic with the posterior
testis reflexed posteriad; situated ventral to the intes-
tine. Spicules strongly curved and bow-shaped, with-
out median stiffening piece. Gubernaculum slightly
curved closer to the dorsal side of the body. Four ejac-

hydr3817.tex; 12/05/1997; 13:18; v.7; p.12

 161

Figure 5. B. fesehai n. sp., male, A & B: Median and surface views respectively of the anterior end; C: Surface view of pharyngo-intestinal
junction; D–E; Tail and reproductive system till the first micropapilla; F: Entire male; G: Testes; H: Pharyngeal region.

ulatory glands present on each side; their ducts clear cells variable in appearance from one part to the oth-
until they reach the level of the anterior end of the er: more granular closer to the cloaca, cells are bigger
spicules, their common duct inconspicuous. Vas defer- and hyaline just posterior to the sphincter, and are
ens long, extending anterior to mid-body; component filled with finer granules anterior to the sphincter. The

hydr3817.tex; 12/05/1997; 13:18; v.7; p.13

162

Figure 6. SEM pictures, B. fesehai n.sp., male, A: Head-on view; B: Dorsolateral view; C: Ventral view showing supplements and the cloacal
opening; D: Inner layer of cuticle of the ventral, posterior body showing transverse striations, laterally observed under LM, these bands appear
as micropapillae; E: A ventromedian supplement. (Scale bar A & D–E = 1 m; B–C = 10 m).

sphincter is short and faintly demarcated, situated far
anterior to the level of the first supplement. Sperma-
tozoa more or less globular each with a dark, usually
centrally positioned spot (nucleus?).
Ventral supplements of two types: six prominent,
bulb-shaped, longer than wide (9.0–10.0 m long and
6.0–8.0 m wide) supplements and up to about 140
micropapillae situated between the bulb-shaped sup-
plements and anterior to the first and posterior to the
last supplement. Each bulb-shaped supplement has 2–3
internal canals that form a common duct leading to the
anterior wall of the ampulla. Distance between bulb-
shaped supplements variable, the shortest distance in
most specimens (23.0–31.0 m) between the fourth
and fifth supplements while the longest distance in
most specimens (27.0–53.0 m) was between the fifth
and sixth supplements (Table 3). Some micropapillae
two-humped. Number of micropapillae variable: 52–
67 anterior to the first supplement, 10–15 between the
first and second supplement, 10–13 between the sec-
ond and third supplement, 11–13 between the third and
fourth supplement, 11–16 between the fourth and fifth
supplement, 19–31 between the fifth and sixth supple-
ment, and 10–14 between the sixth supplement and the
cloacal opening.
Tail conoid in its anterior half; drastically narrow-
ing at about half its length; narrow, cylindrical in its
posterior half; narrow part about 18.0% of cloacal body
width. Six to 8 setae on each side of tail, one terminal
and one sub-terminal seta present.
Population from Zegie
Except for the slightly more anteriorly situated excre-
tory pore, the male specimens from Zegie are similar
to those of the type population.
Diagnosis and relationships
Though the sixth supplement in our new species is
not situated very close to the cloacal opening, it still
is located less than a spicule length from the cloa-
cal opening. Hence, B. fesehai n.sp. falls into Tsalo-
likhin’s (1992) species group ‘B’ (see discussion on

hydr3817.tex; 12/05/1997; 13:18; v.7; p.14

 163

B. graciloides above). Among the species that fall and only six micropapillae anterior to the first supple-
under this category, only B. granatensis (Ocaña & ment in males of B. kenyensis) (Tsalolikhin, 1992).
Zullini, 1988) Tsalolikhin, 1992, B. kenyensis Tsa-
C. – B. malayanus and B. findeneggi
lolikhin, 1992, and B. siculus (Vinciguerra & Zullini,
B. fesehai n.sp. differs from B. malayanus, apart
1991) Ocaña, Hernández & Martin, 1996 come close to
from the above-mentioned, in having narrower anterior
B. fesehai n.sp. in having spicules that are above 40 m
end and less anteriorly situated amphids (LRW = 21–
long. In having shorter cephalic setae B. malayanus
26 m and amphids are situated at the level of the base
(Schneider, 1937) Tsalolikhin, 1983, and B. findeneg-
of the cup-shaped stoma, anterior to the first pouch
gi (Schiemer, 1971) Tsalolikhin, 1983 also come close
in B. malayanus) (Schneider, 1937). Our new species
to B. fesehai n.sp. However, B. fesehai n. sp. differs
also differs from B. findeneggi, apart from the above-
from the first two in being smaller and in having short-
mentioned, in having a higher value for the ratio‘c0 ’,
er outer labial setae and a shorter distance between the
bow-shaped spicules, higher number of micropapil-
two denticles (L = 1400–2400 m and 1321–2190 m;
lae between the sixth supplement and cloacal open-
longest cephalic setae = 6.0–7.0 m and 6.0 m; dis-
ing, relatively far more anteriorly situated sixth supple-
tance between the first and second denticle = 10–14 m
ment and supplements whose duct is directed anteriad
and 10 m, in B. granatensis and B. kenyensis, respec-
(c0 = 2.6–3.0, spicules more or less straight, a single
tively). Our new species also differs from B. malayanus
micropapillae between sixth supplement and cloacal
and B. findeneggi in having longer spicules (spicule
opening, distance between the sixth supplement and
L = 28–32 m and 26 m in B. malayanus and B. find-
cloacal opening = 13–15 m, duct of each supplement
eneggi, respectively).
straight merging with the dorsal side of the supplement
A. – B. granatensis in B. findeneggi) (Schiemer, 1971).
Females of B. fesehai n.sp. differ from those of
D. – B. siculus
B. granatensis in having amphids situated less ante-
Our new species differs from B. siculus in hav-
riorly and shorter tail (Amph-ABE = 8–13 m, tail
ing narrower anterior end, shorter outer labial and
L = 210–280 m in B. granatensis).
cephalic setae, shorter distance between the two teeth,
Males of B. fesehai n.sp. differ from those of
shorter distance between the first and second supple-
B. granatensis in having bow-shaped and thicker
spicules (6.0 at thickest point), supplements of uni-
ment and third and fourth supplement, and longer dis-
tance between the sixth supplement and cloacal open-
form size and closely spaced, numerous micropapillae,
ing, and more numerous micropapillae posterior to the
the fourth and fifth supplements widely spaced, the last
fourth bulb-shaped supplement especially between the
supplement far from cloacal opening, terminal and sub-
last supplement and the cloacal opening (LRW = 26–
terminal setae (spicules only slightly curved, spicule
30 m, Outer LSL = 6–7 m, CSL = 4–5 m, dis-
W as measured from two paratype males = 4.0 m
tance between the two teeth = 10–13 m, distance
at thickest point, the fifth and sixth supplements
between supplements second-third = 33–51 m, third-
smaller than rest, minimum mean distance between
fourth = 59–81 m, sixth-cloacal opening = 15–20 m,
supplements = 35 m, last supplement situated 13–
micropapillae between fourth-fifth supplement = 9–10,
21 m from cloacal opening, terminal seta absent in
fifth-sixth = 10–12 and sixth-cloacal opening = 2 in
B. granatensis) (Ocaña & Zullini, 1988). These differ-
B. siculus).
ences were confirmed by observing paratypes on slide
B. fesehai n.sp., therefore, can be recognized by
N 836 (collection of the University of Ghent).
a combination of characteristics i.e. in having a nar-
B. – B. kenyensis row anterior end, short cephalic and outer labial setae,
Both sexes of B. fesehai n.sp. differ from those of long and bow-shaped spicules, the sixth supplement at
B. kenyensis, apart from the above-mentioned, in hav- a distance of less than a spicule length from cloacal
ing longer tail. Moreover males differ in being thicker opening, numerous micropapillae, and terminal and
and in having a narrower anterior end, lower value subterminal setae. Moreover, amphid size showed sex-
for the ratio‘c0 ’, numerous micropapillae, terminal and ual dimorphism, being larger in males.
subterminal setae (tail L = 201 for male, 240–280 for
female; a = 51.4, c0 = 6.0, LRW = 21.0 m, lack subter-
minal and terminal setae, number of micropapillae= 43

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164

Table 4. Measurements of males and females of Brevitobrilus tsalolikhini n. sp. and females of Epitobrilus setosus (Altherr, 1963)
Tsalolikhin, 1981 from Gedero and Zegie, Lake Tana.

B. tsalolikhini n. sp. E. setosus

Type population from Gedero Zegie Gedero Zegie
Paratype Paratype Holotype
females males male males female females
(n = 2) (n = 2) (n = 2) (n = 1) (n = 7)

L 904.0, 1560.0 1237.0, 1280.0 1330.0 1256.0, 1272.0 1179 1480.3  83.7 (1353.0–1594.0)
LRW 26.0, 28.0 20.0, 21.0 21.0 20.0, 20.0 22.0 26.6  0.9 (25.0–28.0)
Amph–ABE 16.0, 19.0 14.0, 15.0 16.0 15.0, 15.0 23.0 27.9  2.8 (23.0–33.0)
Amph W 5.0, 5.0 6.0, 6.0 6.0 5.5, 5.5 5.0 5.0  0.3 (4.5–5.5)
CBW 27.0, 30.0 24.0, 24.0 24.0 22.0, 22.5 25.0 32.4  1.7 (30.0–34.0)
Stoma L 17.0, 18.0 15.0, 16.0 17.0 14.0, 14.0 20.0 26.5  1.1 (25.0–28.0)
Stoma W 12.0, 13.0 8.5, 8.5 9.0 8.5, 8.5 9.0 12.4  0.9 (11.0–14.0)
Outer LSL 10.0, 11.0 8.0, 8.0 9.0 8.0, 8.0 12.0 13.6  0.9 (12.0–15.0)
CSL 5.0, 7.0 5.0, 5.0 6.0 5.0, 5.0 9.0 5.7  0.5 (5.0–6.0)
First tooth–ABE 22.0, 23.0 18.0, 20.0 20.0 20.0, 20.5 19.0 22.3  1.8 (20.0–24.5)
First–second tooth 10.0, 10.0 10.0, 10.0 11.0 8.5, 10.0 12.0 15.0  1.4 (13.0–17.0)
NR 92.0, 104.0 84.0, 90.0 105.0 90.0, 92.0 94.0 122.1  9.2 (113.0–137.0)
Ph L 202.0, 290.0 213.0, 228.0 270.0 233.0, 246.0 312.0 377.9  12.4 (355.0–392.0)
MBW 60.0, 80.0 56.0, 62.0 70.0 52.5, 52.5 55.0 62.4  5.9 (54.0–70.0)
ABW 30.0, 32.0 27.0, 30.0 31.0 26.0, 27.0 28.0 33.4  2.5 (30.0–38.0)
RL 26.0, 28.0 28.0 34.0  2.8 (28.0–38.0)
Tail L 163.0, 173.0 111.0, 111.0 120.0 109.0, 121.0 170.0 200.0  13.8 (185.0–224.0)
G1L 181.0, 362.0 80.0 150.3  29.3 (105.0–193.0)
G2L 176.0, 321.0 80.0 155.4  28.8 (92.0–179.0)
V 471.0, 751.0 573.0 706.0  48.0 (628.0–775.0)
Egg L 90.7  6.8 (84.0–100.0)
Egg W 40.7  5.6 (33.0–46.0)
Spicule L 31.0, 33.0 35.0 35.0, 36.0
Gubernaculum L 11.0, 13.0 14.0 12.0, 15.0
Distance between supplements
First–second 33.0, 35.0 38.0 33.0, 34.0
Second–third 35.0, 37.0 42.0 37.0, 38.0
Third–fourth 50.0, 50.0 52.0 48.0, 48.0
Fourth–fifth 27.0, 30.0 35.0 28.0, 28.0
Fifth–sixth 40.0, 45.0 50.0 35.0, 35.0
Sixth–cloaca 14.0, 15.0 15.0 12.0, 12.0

a 15.1, 19.5 18.2, 21.4 23.8  1.5 (21.6–25.3)

b 4.5, 5.4 4.9, 5.7 6.0 5.2, 5.4 3.8 3.9  0.2 (3.7–4.1)
c 5.5, 9.0 10.7, 11.1 11.1 10.4, 11.7 6.9 7.4  0.3 (7.1–7.8)
c0 5.4, 5.4 3.9, 4.0 4.1 4.0, 4.7 6.1 6.0  0.4 (5.6–6.9)
V% 48.1, 52.1 48.6 47.7  1.6 (45.2–49.8)
NR% 35.9, 45.5 38.9, 39.4 39.5 38.0, 38.6 30.1 32.3  2.0 (29.8–35.5)
Amph–ABE/LRW 0.6, 0.7 0.7, 0.7 0.8 0.8, 0.8 1.0 1.1  0.1 (0.8–1.3)
Amph W% CBW 16.7, 18.5 25.0, 25.0 25.0 24.4, 25.0 20.0 15.4  0.9 (14.1–16.7)

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3. Brevitobrilus tsalolikhini n.sp.
Measurements
Table 4 (Figures 7, 8 & 9A–F)
Type habitat and locality
Three males and 2 females from surface to 3.0 cm of
sediment at 0.5 m depth, Gedero, L. Tana.
Additional collection
Two males from surface to 1.0 cm sediment at 0.5 m
depth, Zegie, L. Tana.
Type specimens
Holotype male (N 3918) with 2 female and 3 male
paratypes deposited in the collection of the University
of Gent, Instituut voor Dierkunde, Gent, Belgium
Etymology
The species is named after Prof. Dr S. J. Tsalolikhin
in recognition of his contribution to the understand-
ing of the morphology and systematics of the family
Tobrilidae.
Description
Type population
Female:
Body after fixation undulated, tapering towards the
posterior end; maximum body width anterior, at or
sightly posterior to vulva. Cuticle 2.0–3.0 m thick,
smooth under LM; its layers conspicuous, a thinner,
lighter outer and slightly thicker and darker inner lay-
er. Somatic setae few. Below the cuticle is found a thin
layer of fine, dark-brown refractive particles. Eight to
nine dorsolateral, type-II loxometanemes on each side
of the body; the first situated close to the level of the
nerve ring and the last in the tail; length of frontal
and caudal filament about 60 and 65 m, respective-
ly. Metanemes overlapping in most part of the body,
overlap involving only two consecutive metanemes,
the caudal filament of the anterior metaneme and the
frontal filament of following metaneme. Crystalloid
bodies all over the body, situated most probably in the
pseudoceolom.
165
Lip bluntly truncate, continuous with the rest of
the body. Sensilla in two circles: an inner ring of six
labial papillae and an outer ring of four shorter (5.0
& 7.0 m) cephalic and six distinctly longer (10.0 &
11.0 m) labial setae. Amphids stirrup- shaped with
transverse oval aperture, 16.7 & 18.5% CBW; situat-
ed 0.6 & 0.7 LRW from ABE; fusus amphidialis just
behind fovea, as long as fovea. Cheilostome distinct,
with weakly sclerotized lining. Rest of stoma consist-
ing of three parts, an anterior, cup-shaped part followed
by two smaller ventrosublateral pouches of which the
anterior one is situated on the right and the posterior
one on the left side. Pouch wall well sclerotized; each
pouch is supplied with a well sclerotized tooth. Cup-
shaped part 1.3 & 1.5 times longer than wide. First
tooth situated 22.0 & 23.0 m from ABE, the second
tooth situated 10.0 m from the first.
Pharynx cylindrical and muscular. Nerve ring
prominent, at 35.9 & 45.5% pharyngeal length. Car-
dia 17–18  8–19 m. Three prominent (11–15  16–
20 m) cells situated on the junction of the cardia
with the pharynx base. Two highly granular coelomo-
cytes(?) present at the level of these cells or slightly
anterior; coelomocytes longer than wide. Excretory
pore situated about 95.0 m from the ABE, almost
at the level of the nerve ring. Ventral gland cells not
observed. Cells of the intestinal wall filled with brown
granules. Intestine leading to 26.0 & 28.0 m long
rectum.
Reproductive system didelphic, amphidelphic.
Gonoducts situated ventral to the intestine. Spermath-
eca indistinct but spermatozoa observed in uterus. The
latter supplied with well developed, externally trans-
versely striated muscular wall. Ovejector with wide
lumen; its wall internally lined by a highly plicated
layer. Vagina about 14.0% CBW; sphincter muscle
inconspicuous. Vaginal wall without plication. Vul-
va slightly anterior or slightly posterior to mid-body.
Up to 4 eggs present in each uterus.
Tail elongate-conoid, about 6.0 rectal lengths long,
straight, gradually narrowing for most part with the last
about 10 m long posterior end being slightly swollen
and club-shaped. Nine to ten setae on tail; subterminal
seta present, terminal seta absent. Caudal glands three,
prominent; terminating in a short, tubular spinneret
outlet that opens perfectly terminal on the otherwise
blunt tail end.
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166

Figure 7. B. tsalolikhini n.sp., female, A: Reproductive system; B: Pharyngeal region till the nerve ring and excretory pore; C & D: Median
and surface views respectively of rectal region and tail; E: Surface view of anterior end; F: Entire female.

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 167

Figure 8. B. tsalolikhini n.sp., male, A & D: Median and surface views respectively of the anterior end; B–C: Tail and reproductive system till
the anterior sphincter; E: Pharyngeal region; F: Surface view of pharyngo-intestinal junction; G: Surface view of tail; H: Testes; I: Entire male.

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168

Figure 9. SEM pictures, A–F B. tsalolikhini n.sp., male, A: Head-on view; B: Ventral view of the body between the last two ventromedian
supplements showing the two longitudinal, ventrosublateral, cuticular grooves (arrowheads); C: A ventromedian supplement; D: The last
ventromedian supplement, transverse cuticular incisures (appear as micropapillae when seen laterally under LM), and cloacal opening; E:
Longitudinal section of the anterior end showing cup-shaped part of the stoma and the position of dorsal gland opening (arrow); F: Dorsal gland
opening (arrow); G: Longitudinal section of the anterior end of E. setosus showing the anterior funnel-part, the posterior, left, ventrosublateral
pouch and its associated tooth. (Scale bar A–B, D–E & G = 10 m; C & F = 1 m).

Type population
Male:
Resembling female in many respects except the
following (only males were studied under the scanning
electron microscope). Body after fixation curved dor-
sad; maximum body width often at the level of the
testes. Cuticle faintly striated under SEM. Amphids
wider than in females, 25.0% CBW. Dorsal gland open-
ing a longitudinal slit, situated posterior to the level of
the first ventrosublateral pouch (Figure 9F).
Reproductive system diorchic with the posterior
testis reflexed posteriad; situated ventral to the intes-
tine. Spicules only slightly curved, without median
stiffening piece. Gubernaculum straight. Six ejacula-

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tory glands present on each side; their ducts merging
together to form a common duct ventral to the anterior
end of each spicule. Vas deferens long, extending ante-
rior to mid-body; component cells variable in appear-
ance from one part to the other: hyaline close to the
cloaca until the level of the anterior sphincter, filled
with fine granules more anteriorly. Two sets of sphinc-
ters observed, the first sphincter muscles are situated
about 55 m anterior to the first supplement while the
second set is an equal distance anterior to the first set.
Spermatozoa elongated, 10–17 m long, wider on one
end and tapering on the other, up to 3.5 m wide at its
widest part.
SEM pictures showed two longitudinal, subven-
tral, cuticular grooves, one right and the other left,
extending from the cloacal opening to the level of the
most anterior supplement (Figure 9B). Ventral sup-
plements of two types: six prominent, laterally bulb-
shaped, longer than wide (8.0–14.0 m long and 8.0–
11.0 m wide) supplements and up to about 46, lateral-
ly observed, papilloid supplements between the bulb-
shaped supplements and anterior to the first and poste-
rior to the last supplement. Each bulb-shaped supple-
ment has 2–3(?) internal canals forming a straight com-
mon duct that merges with the dorsal side of the ampul-
la. Under SEM these supplements externally appear to
consist of two concentric rings with a central plug-like
structure. Distance between bulb-shaped supplements
variable, the shortest distance (27.0, 30.0 & 35.0 m)
was between the fourth and fifth supplement while the
longest distance (50.0, 50.0 & 52.0 m) was between
the third and fourth supplement (Table 4). Under SEM
micropapillae appear to be ventral bands, about 12–
15 m wide and 2–3 m long, each separated from the
other by a transverse groove (incisure); their number
variable: 8, 8 & 9 anterior to the first supplement, 6
between the first and second supplement, 6, 7, & 8
between the second and third supplement, 9, 9 & 10
between the third and fourth supplement, 6 between
the fourth and fifth supplement, 7, 8 & 8 between the
fifth and sixth supplement, and 1, 2 & 2 between the
sixth supplement and cloacal opening.
Tail elongate-conoid in its anterior half, narrow
and cylindrical more posteriorly; narrowest part about
13.0% of cloacal body width. Ten to eleven setae on
each side of tail, one subterminal seta present at the
5–6 m from PBE.
169
Specimens from Zegie
Except for the slightly narrower anterior end, the male
specimens from Zegie are similar to those of the type
population.
Diagnosis and relationships
The sixth supplement in B. tsalolikhini n.sp. is situated
very close to the cloacal opening (Table 4). Compared
with those species that have their sixth supplement at
a distance less than a spicule length from the cloacal
opening, our new species comes close to B. findeneggi,
B. granatensis, B. kenyensis and B. malayanus. How-
ever, B. tsalolikhini n.sp. differs from all four species in
its longer cephalic and outer labial setae (CSL = 2.6 m
for male, 2.9 m for female; outer LSL = 2.9 m
for male, 3.2 m for female in B. findeneggi; longer
cephalic setae = 6–7 m in B. granatensis; 6.0 m in
B. kenyensis; and 4.0 m in B. malayanus). B. tsalo-
likhini n.sp. differs from the last three species also in
having supplements whose straight duct merges with
the dorsal wall of the ampulla (duct of supplements
directed anteriad and merging with anterior wall in
B. granatensis, in B. kenyensis and in B. malayanus).
B. tsalolikhini n.sp. can easily be differentiated from
all known species in the genus in having a combina-
tion of characteristics: long cephalic and outer labial
setae, short spicules without median stiffening piece,
the sixth supplement situated very close to the cloacal
opening, supplements with straight duct that merges
with the dorsal wall of the ampulla, and elongated
spermatozoa.
A. – B. findeneggi
Compared to our new species, B. findeneggi has
narrower stoma, shorter spicules, higher value for
the ratio‘c’, and lower value for the ratio‘c0 ’ (stoma
W = 5.0 m for female, 6.0 m for male; spicule
L = 26.0 m, c = 13.1 for female, 18.5 for male; c0 = 2.6
for male, 3.0 for female, in B. findeneggi) (Schiemer,
1971).
B. – B. granatensis
B. tsalolikhini n.sp. differs from B. granatensis,
apart from the above-mentioned, in having: shorter
tail, relatively prominent micropapillae, and elongated
spermatozoa (tail L = 210 m in female, 126–165 m
for male, micropapillae inconspicuous, and spermato-
zoa more or less globular in B. granatensis) (Ocaña &
Zullini, 1988). Besides, our examination of the type
specimens of B. granatensis has confirmed these dif-
ferences.
hydr3817.tex; 12/05/1997; 13:18; v.7; p.21
170
C. – B. kenyensis
Our new species differs from B. kenyensis, apart
from the above-mentioned, in being thicker and in
having shorter spicules, its sixth supplement slightly
closer to the cloacal opening, and shorter tail (a = 22.4–
26.7 for female, 51.4 for male, spicule L = 48.0 m,
distance of the sixth supplement from cloacal opening
20.0 m, tail L = 201 for male, 240–280 m for female
in B. kenyensis) (Tsalolikhin, 1992).
D. – B. malayanus
B. tsalolikhini n.sp. differs from B. malayanus,
apart from the above-mentioned, in having its sixth
supplement closer to the cloaca and fewer micropapil-
lae between its sixth supplement and cloacal opening
(distance of the sixth supplement from cloacal open-
ing = 21.5–26.4 m, number of micropapillae between
the sixth supplement and cloacal opening = 6–8 in
B. malayanus) (Schneider, 1937).
E. – B. fesehai n.sp.
B. tsalokihini n.sp. differs from B. fesehai n. sp.
in the shape of its anterior end and tail, width of
lip region, length of cephalic and labial setae, length
of cup-shaped part of stoma, shape and length of
spicules, position of attachment of the duct of supple-
ment with ampulla wall, shape of supplement, distance
between supplements, distance of the sixth supplement
from cloacal opening, size, number and distribution of
micropapillae, shape of spermatozoa, in the value for
Amph% CBW, and in lacking a terminal seta.
On the identification of species in the genus
Brevitobrilus
Almost every characteristic of both sexes has been used
to differentiate one species from the other in Brevito-
brilus. However, more recently a tendency to use male
copulatory structures for a similar purpose has domi-
nated the descriptions, because male copulatory struc-
tures give more differentiating possibilities among the
species (Ocaña & Zullini, 1988; Tsalolikhin, 1992).
Despite this tendency, the quality of descriptions in the
group remains variable. Riemann (1983), in his brief
review, clearly pointed out the usefulness of the shape
of spermatozoa in the nematodes in general and in the
then genus Tobrilus in particular. Nevertheless, none
of the species descriptions in the genus includes the
shape of spermatozoa. This, we believe, is a serious
drawback in the understanding of the morphology of
the genus. Therefore, we suggest that future descrip-
tions of amphimictic species include description and
illustrations of spermatozoa.
4. Epitobrilus setosus (Altherr, 1963) Tsalolikhin,
1981
Measurements
Table 4 (Figures 9G & 10)
Description
Population from Zegie
Female:
Body after fixation straight, tapering towards both
ends, but more so towards the posterior end; maxi-
mum body width mostly at level of vulva. Cuticle 1.5–
2.0 m thick; its layers faintly conspicuous, a thinner,
lighter outer and slightly thicker and darker inner lay-
er. Somatic setae distributed all over the body. Ten
to twelve dorsolateral type II loxometanemes on each
side of the body; the first situated close to the level of
the nerve ring and the last in the tail; length of frontal
and caudal filament about 95 and 90 m, respectively.
Mostly only two metanemes overlapping, some times
three metanemes overlapping; in the first case, a caudal
filament of anterior metaneme and frontal filament of
following metaneme overlap or entire caudal and part
of frontal filament of an anterior metaneme overlapping
with the entire frontal and part of caudal filament of
following metaneme; in the latter case, part of caudal
filament of a metaneme overlaps with part of the frontal
filament of the next metaneme and part of the frontal
filament of a third metaneme. Crystalloid bodies all
over the body (most probably in the pseudocoelom),
concentrated in the pharyngeal region.
Lip region bluntly round, continuous with the rest
of the body. Sensilla prominent, in two circles: an
inner ring of six labial setiform papillae (1.5–2.0 m
long) and an outer ring of four shorter (5.0–6.0 m)
cephalic and six distinctly longer (12.0–15.0 m)
labial setae. Amphids stirrup-shaped, inconspicuous
in most specimens, with transverse oval aperture,
14.1–16.7% CBW; situated 0.8–1.3 LRW from ABE;
fusus amphidialis just behind fovea, as long as fovea.
Cheilostome distinct, with weakly sclerotized lining.
Rest of stoma consisting of two distinct parts, an ante-
rior, funnel-shaped part consisting of the cup-shaped
part of the stoma and two ventrosublateral pouches
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 171

Figure 10. E. setosus, female, A & F: Dorsoventral and lateral median views respectively of the anterior end; B: Entire female; C: Pharyngeal
region; D: Rectal region and tail; E: Reproductive system; G: Surface view of anterior end; H: Surface view of pharyngo-intestinal junction.

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172
Table 5. Measurements of males, females and intersex of Tripyla glomerans Bastian, 1865 from Zegie, Lake Tana.

Female Male Intersex

(n = 10) (n = 10) (n = 3)

L 2218.0  240.4 (1904.0–2790.0) 2261.5  260.2 (1858.0–2790.0) 1683.0, 2102.0, 2390.0

LRW 43.5  2.2 (41.0–47.0) 41.8  2.0 (39.0–47.0) 43.0, 43.0, 44.0
LRH 28.0  2.8 (26.0–32.0) 29.4  2.9 (27.0–33.0) 27.8, 29.0, 31.0
Amph–ABE 21.3  2.4 (17.0–24.0) 20.4  1.9 (17.0–23.0) 18.0, 21.0, 22.0
Amph W 7.0  0.5 (6.0–8.0) 7.0  0.2 (6.5–7.5) 7.0, 7.0, 7.0
CBW 43.1  3.3 (36.0–47.0) 41.9  2.1 (39.0–47.0) 41.0, 43.0, 44.0
OLSPL 2.5  0.5 (2.0–3.5) 2.5  0.4 (2.0–3.0) 2.0, 3.0, 3.0
CSL 3.9  0.5 (3.5–5.0) 3.9  0.5 (3.0–5.0) 4.0, 4.0, 4.0
Dorsal tooth–
ABE 24.1  1.4 (22.0–26.0) 24.4  1.3 (23.0–27.0) 25.0, 25.0, 25.0
NR 122.2  9.0 (112.0–140.0) 126.0  7.8 (107.0–135.0) 112.0, 130.0, 135.0
Ph L 371.8  36.2 (324.0–441.0) 369.0  21.5 (333.0–400.0) 342.0, 385.0, 389.0
Ex. pore–ABE 74.1  3.8 (68.0–79.0) 75.3  8.1 (62.0–84.0) 72.0
MBW 82.1  6.7 (73.0–93.0) 80.0  4.3 (73.0–86.0) 63.0, 70.0, 84.0
ABW 53.0  4.7 (47.0–60.0) 58.7  4.2 (51.0–66.0) 51.0, 51.0, 54.0
RL 44.4  3.2 (41.0–51.0)
Tail L 419.1  33.1 (378.0–491.0) 436.7  41.5 (369.0–508.0) 380.0, 396.0, 455.0

G1L 322.3  93.4 (238.0–522.0) 252.0, 270.0

G2L 334.4  123.4 (234.0–617.0) 234.0, 252.0
V 1095.8  144.6 (856.0–1386.0) 828.0, 1088.0, 1218.0

Spicule L 69.5  3.1 (65.0–75.0) 33.0–43.0, 54.0, 59.0

Gubernaculum L 20.7  1.1 (19.0–23.0)
Total number of supplements 12.0  2.5 (9.0–16.0)
Number of supplements on neck 6.0  0.9 (5.0–7.0)
a 27.1  2.8 (23.5–32.4) 28.3  3.1 (23.8–33.2) 24.0, 28.5, 33.4
b 6.0  0.3 (5.5–6.3) 6.1  0.7 (5.4–7.8) 4.9, 5.4, 6.2
c 5.3  0.5 (4.6–5.9) 5.2  0.9 (4.3–6.0) 4.4, 5.3, 5.3
c0 8.1  1.0 (6.3–9.1) 7.5  0.9 (5.6–8.6) 7.3, 7.5, 8.9
V% 49.3  2.2 (45.0–51.2) 49.2, 51.0, 51.7
NR% 33.4  1.7 (31.7–36.3) 34.2  2.0 (31.3–38.4) 32.7, 33.4, 35.1
Amph–ABE/LRW 0.5  0.1 (0.4–0.6) 0.5  0.1 (0.4–0.6) 0.4, 0.5, 0.5
Amph% CBW 16.4  2.0 (14.3–22.20) 16.9  1.3 (14.9–17.9) 15.9, 16.3, 17.1

with well sclerotized wall, one on the right followed
by the second on the left. Funnel-shaped part 1.9–2.3
times longer than wide. First tooth situated 20-24.5 m
from ABE, second tooth 13.0–17.0 m posterior to the
first.
Pharynx cylindrical for most part of its length,
slightly narrower at its posterior end; muscular. Nerve
ring prominent, at about a third of pharyngeal length.
Cardia 19–20  20-22 m. Three prominent (15-
22  18–25 m) cells situated at the junction of the car-
dia with the pharynx base. A coelomocyte (?) present
anterior to these cells; coelomocytes longer than wide.
Excretory pore situated about 140.0 m from the ABE,
at the level of the nerve ring. Ventral gland cells not
observed. Cells of intestinal wall filled with brown
granules. Intestine leading to 28.0–38.0 m long rec-
tum.
Reproductive system didelphic, amphidelphic.
Gonoducts situated ventral to the intestine. Uterus sep-
arated from the rest by a well developed sphincter.
Vagina 19.0–23.0% CBW; with well developed sphinc-

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 173

ter muscle. Vulva slightly anterior to mid-body. Only
one egg at a time in uterus; eggs longer than wide.
Tail elongate-conoid, gradually tapering till about
20 m from PBE, slightly swollen and club-shaped
more posteriorly; straight; about 6.0 rectal lengths
long. Two to five setae on tail; subterminal and ter-
minal seta absent. Caudal glands three, prominent;
terminating in a short, tubular spinneret outlet that in
some specimens is situated perfectly terminal on the
blunt tail end.
Male not found.
azii (Eyualem & Coomans, 1996). Morphometrical-
ly, our population differs from the original population
of E. setosus in its size and values for the ratios‘a’
and ‘c’ (L = 2190–2210 m, a = 30.0–33.0, c = 8.3–9.1
in the original population of E. setosus). In spite of
the absence of long ‘setae’ and the obviously smaller
size in our population, due to the similarities in other
respects, we prefer to consider them as a population of
E. setosus.
This is the first report of E. setosus out of its type
locality, Argentina.

Comparisons and rationale for identification

5. Tripyla glomerans Bastian, 1865
Tsalolikhin (1981) listed four species in the genus
Epitobrilus Tsalolikhin, 1981; these were E. flagel-
Measurements
latus (Andrássy, 1963) Tsalolikhin, 1981, E. meyli
Tsalolikhin, 1981, E. parvipapillatus (Kreis, 1932)
Table 5 (Figures 11, 12 & 13)
Tsalolikhin, 1981, and E. setosus (Altherr, 1963)
Tsalolikhin, 1981. More recently, he reassessed the
Description
species in the genus and regarded E. parvipapilla-
tus as species incertae sedis (Tsalolikhin, 1991) and
Female:
transferred Eutobrilus medius (Schneider, 1916) Tsa-
Body after fixation in most cases strongly curved
lolikhin, 1981 to the genus Epitobrilus.
ventrad, rarely undulated; tapering mainly towards the
Comparisons of our population with the four
posterior end; maximum body width anterior, at or
species show that it comes closer to E. medius and
slightly posterior to vulva. Cuticle with the appearance
E. setosus and clearly differs from the other two.
of two detached layers; its thickness variable: 4.0–
E. flagellatus is a smaller species characterized by a
5.0 m at lip region, 8.0–12.0 m around mid-body,
uniquely long tail. The vulva is also more anterior
and 2.0–3.0 m at the posterior part of tail. Both lay-
than in our population (L = 960.0–1040.0 m, c = 3.4–
ers of cuticle annulated; the outer layer with widely
3.9, V% = 43.0–45.2 in E. flagellatus). E. meyli is also
spaced annuli (annulus W = 2.5–3.0 m), inner layer
unique in the genus in having its first tooth in the middle
with narrower annuli (annulus W = 1.0–1.5 m). First
of the exceptionally broad anterior stoma; it also has
annulus situated 26.0–32.0 m from ABE. Annuli of
shorter outer labial setae (outer LSL = 9.0–10.0 m).
the outer layer inconspicuous on extreme posterior part
The differences of our population with E. medius are its
of tail. Somatic setae absent. Scarcely distributed body
smaller size and lower value for the ratio ‘c’ (L = 2500–
pores present. Each body pore supplied with an internal
3300 m, and c = 10–12 for female in E. medius).
plug-like structure (seta?) that was never seen protrud-
Moreover, E. medius, as shown in the original illus-
ing out of the body pore. Their distribution on each side
trations, is characterized by having a terminal seta
of the body as follows: 4–5 subdorsal and 4 subventral
(Schneider, 1916). Tsalolikhin (1991) denoted this seta
on pharyngeal region, 3–4 subdorsal and 2–3 subven-
in this species as subterminal. In most morphological
tral between base of pharynx and vulva, 1–2 subdorsal
characteristics, our species is similar to E. setosus, the
and 2 subventral between vulva and anus, and 2 lateral
only difference being the absence of long (12–15 m)
on tail. Metanemes inconspicuous in most specimens,
somatic setae distributed all over the body. Somatic
seen only in three specimens, a maximum of three
setae in our population are short (3.0–3.5 m). But,
orthometanemes on each side of the body. Very dense-
similar to our population, the absence of terminal and
ly packed crystalloid bodies present all over the body,
subterminal setae was clearly stated in the original
situated in the pseudocoelom.
description (Altherr, 1963). Though we cannot be sure
Lip region bluntly rounded, slightly narrower than
what was depicted as somatic setae by Altherr (1963);
the adjacent body; smooth. Lips united to form three
long setae-like foreign particles that could be strings of
lobes, the two subdorsal lips forming the dorsal lobe,
bacteria have been observed in Eumonhystera mwer-
each lateral lip with each of the subventral lip forming

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174

Figure 11. T. glomerans, A–I female, A & B: Surface and median views respectively of the anterior end; C: Anterior branch of reproductive
system of a mature female; D & E: Surface and median views of pharyngo-intestinal junction; F: Reproductive system of a young female; G:
Entire female; H: Rectal region and tail; I: Tail tip and spinneret; J–L intersex, J: Female reproductive system of an intersex; K & L: Male
copulatory structures of an intersex showing two small spicules on one side of the body (K), and single spicule on one side of the body (L).

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Figure 12. T. glomerans, male, A: Male copulatory structures; B: Pharyngeal region till nerve ring; C: Testes; D: Surface view of anterior end;
E: Tail and posterior part of male reproductive system; F: Spermatozoa; G: Entire male; H & I: Median and surface views respectively of the
pharyngo-intestinal junction; J: tail tip and spinneret.
the two ventrolateral lobes, giving a triradiate appear-
ance to the mouth opening. Internally two incompletely
separated liplets stick out from each lobe. Sensilla in
three circles: an inner ring of six labial papillae sit-
uated at the border of the mouth opening, a second
ring of six outer labial setiform papillae (2.0–3.5 m)
and a more posterior ring of four slightly longer (3.5–
4.0 m) cephalic setae. Amphids stirrup-shaped with
transverse oval opening, situated in the unstriated part
of the anterior end, 14.3–22.2% CBW; situated 0.4–0.6
LRW from ABE; fusus amphidialis just behind fovea,
as long as fovea. Stoma narrow and indistinct. A pair
of minute ventrosublateral denticles and a single rel-
atively prominent dorsal tooth present, dorsal tooth
situated 22.0–26.0 m from ABE. The ventrosublater-
175
al denticles situated 2.5–3.5 m anterior to the dorsal
tooth. Dorsal pharyngeal gland opening close to base
of (through?) the dorsal tooth.
Pharynx cylindrical and muscular. Nerve ring at
about one-third of pharyngeal length. Cardia as wide
as long. Three prominent (12  20–22 m) cells are
situated at the junction of the cardia with the pharynx
base. On each side of the body, close to the base of
the pharynx is situated a prominent coelomocyte (?).
Coelomocyte wider than long. Excretory pore situat-
ed 68.0–79.0 m from the ABE. Ventral gland cells
not observed. Cells of intestinal wall filled with brown
granules. Intestine leading to 41.0–51.0 m long rec-
tum.
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176

Figure 13. SEM pictures, T. glomerans, A–D female, A: Lateral view of anterior end showing outer labial setiform papillae (white arrows) and
cephalic setae (black arrows); B & C; Head-on views showing inner labial papillae (arrowheads), outer labial setiform papillae (white arrows)
and cephalic setae (black arrows); D: Tail tip and spinneret opening; E: Ventromedian supplement (arrow) in male. (Scale bar A–C = 10 m;
D–E = 1 m).

Reproductive system didelphic, amphidelphic. Male:

Position with respect to intestine variable. Spermath-
eca indistinct in young females, distinct and offset
in mature females, full of congested spermatozoa in
impregnated females. Vagina with thick and well cutic-
ularized wall, 44.5–50.6% CBW; with well developed
sphincter muscle. Vulva anterior to or at mid-body.
Tail elongate-conoid gradually narrowing posteri-
orly, about 9 rectal lengths long, usually curved ven-
trad. Caudal glands three, granular; two of them almost
at the same level, close to the rectum, the third situated
more posteriorly; terminating in about 11.0 m long
spinneret. Spinneret opening terminal.
Resembling female in many respects except the
following. Posterior part of body more strongly curved
ventrad than is in female; maximum body width often
at the level where the two testes join the gonoduct. Dis-
tribution of pores on each side of the body as follows: 4
subdorsal and 4 subventral on pharyngeal region, 6–7
subdorsal and 2–4 subventral between base of pharynx
and cloacal opening, and 2 lateral on tail. Three gran-
ular coelomocytes present anterior to the level of the
cardia.
Reproductive system diorchic with the posterior
testis reflexed posteriad; situated ventral to the intes-
tine. Spicules horn-shaped, wider anteriorly and grad-

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ually narrowing and slightly curved ventrad towards
their tip. Their wider part entangled in a well developed
muscular cap and dorsally extending retractor muscle.
Gubernaculum straight and short with a hook-shaped
distal side. A pair of, one short and one long, ejac-
ulatory glands present on each side of the body. Vas
deferens long, extending anterior to mid-body; compo-
nent cells uniformly granular. Spermatozoa elongated,
13.0–20.0 m long, about 5.0 m wide at the widest
end, tapering on one end.
Ventral supplements in the form of small papillae,
distributed along the ventromedian line, the first situ-
ated anterior to the excretory pore, 55.0–65.0 m from
ABE; the last situated 60.0–70.0 m anterior to the
cloacal opening.
Intersex (n = 3):
Resembling female in every respect except the fol-
lowing. In addition to female characteristics, these
three specimens had male characteristics i.e. possessed
spicules and a single ventromedian supplement. Two
of the three intersex specimens possessed a pair of
spicules; each spicule attached to a well developed
retractor muscle. In these specimens the single ven-
tromedian papilloid supplement was situated about
75.0 m from the cloacal opening. The third specimen
possessed three spicules, a pair of smaller spicules on
the right and a single, bigger spicule on the left side.
In this specimen the only ventromedian supplement
was situated more posteriorly (45.0 m from cloacal
opening). Gubernaculum, vas deferens and testis were
absent. No spermatozoa observed in the female repro-
ductive system.
Comparisons and rationale for identification
Brzeski & Winiszewska-Slipinska (1993), though indi-
cated the number of species in the genus Tripyla Bas-
tian, 1865 to be 16, they actually listed 17 species,
among which three were new. Compared to the 17
species, our population comes close to only one
species, T. glomerans Bastian, 1865. Same authors
stated that ‘T. glomerans can be recognized because
of large body, cephalic sensilla larger than outer labi-
al ones, body pores along lateral chord, tail shape,
spicule and gubernaculum length and shape, and large
number of ventromedian supplementary papillae’. In
all morphological and most morphometrical character-
istics our population is similar to T. glomerans. The
only morphometrical differences are the slightly more
anteriorly situated excretory pore and vulva (excretory
pore 94.0–103.0 m from ABE and V% = 51.0–60.0
177
in Polish populations of T. glomerans). Therefore we
have identified our population as such.
Filipjev (1931) was the first to report this species
from Ethiopia. He found it in Lake Ziquala, a small
crater lake at the top of Mt. Ziquala, south-central
Ethiopia. More recently, Tudorancea & Zullini (1989)
reported it from Lakes Ziway and Langano. However,
none of the above authors gave description or illus-
tration of the populations they reported. This report,
in addition to being the first report of the species in
Ethiopia out of the rift valley, also provides a detailed
account of morphological and morphometrical charac-
teristics of the species.
6. Ironus ignavus Bastian, 1865
Measurements
Table 6. (Figure 14)
Description
Population from Gedero
Female:
Body posture after fixation slightly (one specimen)
to strongly curved dorsad (two specimens). Maximum
body width between base of pharynx and halfway the
distance between vulva and anus. Cuticle smooth; lat-
eral body pores absent. Metanemes inconspicuous in
most, two orthometanemes seen only in one specimen.
Crystalloid bodies distributed in all parts of the body;
probably in the pseudocoelom as well as in the mus-
culature; their size ranges from 1.0–10.0 m wide and
1.0–12.0 m long, and their shape varies from spindle-
shaped to rectangular.
An inner circle of six outer labial papillae and an
outer circle of four cephalic setae on the lip region.
Inner labial sensilla inconspicuous, probably papilloid
and small. Outer labial papillae on the anterior side of
lip region. Cephalic setae 22.7–31.8% LRW, situated
on the sides of the lip region, at the level immediate-
ly anterior to the weak constriction-line between the
slightly but detectably bulging lip region and the rest
of the body. Cheilostome short (2.0–2.5 m long) and
narrow. Rest of stoma well sclerotized, very long, and
composed of two parts, an anterior wider chamber fol-
lowed by a more or less cylindrical, posterior part.
Cylindrical part gradually narrows posteriorly. Three
anteriorly curved and strongly developed teeth, one
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 178
Table 6. Measurements of females of Ironus ignavus Bastian, 1865 from Deepest Site and Gedero, L. Tana, and Ironus sphincterus Ebsary, 1985 from
Gedero and Zegie, L. Tana, and L. Ziway.

I. ignavus I. sphincterus
Deepest Gedero Gedero (L. Tana) Zegie (L. Tana) L. Ziway
Site (n = 1) (n = 4) (n = 2) (n = 4) (n = 4)

L 1521.0 1691.8  95.6 (1566.0–1785.0) 1948.0, 3162.0 2824.8  116.8 (2688.0–3004.0) 2801.3  207.2 (2635.0–3156.0)
LRW 11.0 11.0  0.0 (11.0–11.0) 15.0, 15.0 15.5  1.7 (14.0–18.0) 16.0  0.0 (16.0–16.0)
CSL 3.5 3.0  0.4 (2.5–3.5) 4.5, 5.0 4.5  0.4 (4.0–5.0) 4.0  0.6 (3.5–5.0)
Amph W 4.5 5.0  0.0 (5.0–5.0) 7.0, 8.0 9.7  0.5 (9.0–10.0) 7.6  0.2 (7.5–8.0)
CBW 12.0 12.0  0.0 (12.0–12.0) 15.0, 15.0 17.0  1.4 (15.0–18.0) 15.9  0.2 (15.5–16.0)
Stoma L 82.0 80.7  3.1 (78.0–82.0) 103.0, 105.0 99.3  6.9 (95.0–105.0) 97.5  2.3 (95.0–100.0)
Stoma W 4.0 4.3  0.3 (4.0–4.3) 5.0, 6.0 6.4  0.6 (5.5–7.0) 6.0  0.4 (5.5–6.5)
NR 142.0 143.5  5.0 (135.0–148.0) 173.0, 189.0 184.8  9.8 (169.0–195.0) 179.0  0.7 (178.0–180.0)
Ph L 371.0 391.5  5.6 (382.0–396.0) 501.0, 525.0 587.5  9.4 (577.0–602.0) 506.5  15.2 (493.0–532.0)
Cardia L 10.0 10.3  1.6 (9.0–13.0) 17.0, 19.0 14.5  2.6 (10.0–16.0) 18.0  2.9 (14.0–22.0)
Cardia W 12.0 11.0  2.3 (7.0–13.0) 15.0, 22.0 21.5  2.1 (20.0–25.0) 17.5  4.3 (15.0–25.0)
CBW 29.0 35.8  1.5 (34.0–38.0) 51.0, 63.0 63.5  8.3 (52.0–75.0) 50.8  0.4 (50.0–51.0)
G1L 191.0 348.0  55.8 (270.0–426.0) 353.0, 810.0 589.8  106.0 (500.0–762.0) 589.3  63.0 (482.0–644.0)
G2L 210.0 364.3  67.3 (300.0–476.0) 299.0, 850.0 634.8  52.8 (556.0–690.0) 576.5  85.6 (494.0–705.0)
V 792.0 904.5  48.1 (853.0–966.0) 1071.0, 1531.0 1492.8  36.5 (1452.0–1547.0) 1479.0  102.9 (1349.0–1627.0)
Vagina L 13.0 17.7  0.5 (17.0–18.0) 25.0, 33.0 31.3  0.9 (30.0–32.0) 26.5  1.1 (25.0–28.0)
CBW 34.0 35.8  1.5 (34.0–38.0) 52.0, 65.0 66.7  5.0 (60.0–72.0) 50.4  1.1 (49.0–52.0)
MBW 32.0 36.0  1.4 (34.0–38.0) 52.0, 66.0 68.8  5.4 (60.0–75.0) 52.9  0.5 (52.0–53.5)
ABW 17.0 19.8  1.3 (18.0–21.0) 25.0, 30.0 30.0  3.8 (25.0–35.0) 26.1  0.5 (25.5–27.0)
RL 34.0 25.5  6.9 (25.0–27.0) 37.0, 46.0 42.5  3.6 (38.0–47.0) 35.5  2.7 (33.0–40.0)
Tail L 169.0 152.8  14.4 (137.0–176.0) 157.0, 191.0 213.0  5.2 (204.0–216.0) 220.0  21.2 (197.0–245.0)
a 47.5 47.1  3.5 (41.2–49.6) 37.5, 47.9 41.3  2.9 (38.4–46.1) 53.0  3.8 (50.7–59.5)
b 4.1 4.3  0.2 (4.1–4.5) 3.9, 6.0 4.8  0.2 (4.6–5.1) 5.5  0.4 (5.1–6.3)
c 9.0 11.1  0.6 (10.1–11.8) 12.4, 16.6 13.3  0.7 (12.4–13.9) 12.8  1.2 (10.8–13.8)
c0 9.9 7.8  1.2 (6.5–9.8) 6.3, 6.4 7.2  1.0 (6.2–8.6) 8.4  0.7 (7.6–9.4)
V% 52.1 53.5  1.0 (51.9–54.6) 48.4, 55.0 52.9  1.9 (51.1–56.0) 52.8  2.0 (51.2–56.2)
NR% 38.3 36.6  0.8 (35.3–37.7) 30.0, 34.5 31.5  1.9 (28.7–33.8) 35.4  1.1 (33.6–36.5)
Amph% CBW 37.5 41.7  0.0 (41.7–41.7) 46.7, 53.3 57.0  2.1 (55.6–60.0) 48.0  1.3 (46.9–50.0)
CSL% LRW 36.8 27.3  3.2 (22.7–31.8) 30.0, 30.0 29.2  1.7 (27.8–32.1) 25.0  3.8 (21.9–31.3)

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 179

Figure 14. I. ignavus, female, A: Rectal region and tail; B: Entire female; C: Reproductive system; D: Pharyngeal region till nerve ring; E:
Surface view of anterior end; F: Pharyngo-intestinal junction.

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180
simple dorsal and two ventrosublateral, in the wider
part of the stoma. Ventrosublateral teeth slightly larger
than dorsal one. Three pharyngeal gland openings in
the stoma: two anteriorly situated (about 9.0 m from
ABE) ventrosublateral gland openings and more pos-
teriorly (about 35.0 m from ABE) one dorsal gland
opening. Amphids stirrup shaped, opening situated at
the level of lip region constriction, about two-fifths
of corresponding body width wide. Pharynx cylindri-
cal for most of its length somewhat constricted at the
level of the nerve ring. Where exactly the pharynx
musculature starts is not evident, but, muscles become
unambiguously conspicuous on the dorsal side around
the middle of the stoma. On the ventral side, the pha-
ryngeal muscles become conspicuous about 10.0 m
anterior to the end of the stoma. Nerve ring surrounds
pharynx at about a third to two-fifths of pharyngeal
length. Excretory pore and ventral gland not seen. Car-
dia triangular, slightly more than one-third of the corre-
sponding body width. Intestine with distinct microvilli,
its cells filled with brown granules, leading to a 25.0–
27.0 m long rectum.
Female reproductive system didelphic, amphidel-
phic with reflexed ovaries, position of reproductive
system with respect to intestine variable. Uterus sepa-
rated from remainder of gonoduct by a weakly muscu-
lar sphincter. Vagina muscular, about half CBW long.
Vulva a transverse slit, situated slightly posterior to
mid- body. Eggs 5-8 times longer than wide, 20.0–
25.0 m  127–160 m.
Tail gradually tapering to a fine sharp tip; about 6
rectal lengths long.
Male not found.
Specimen from deepest site
This specimen showed similarity in every respect
except for its slightly smaller size, narrower body,
slightly narrower amphid, shorter vagina, and longer
rectum.
Comparisons and rationale for identification
In being small and didelphic, in having stoma less
than 90.0 m, ‘c’ less than 20.0, ‘c0 ’ greater than 5.0,
a vulva situated behind mid-body, and in lacking a
sclerotized, refractive sphincter, our specimens come
close to two species viz. I. americanus Cobb, 1914
and I. ignavus Bastian, 1865. These two species are
similar but, they can be differentiated from each other
by their respective values for the ratios‘a’ and ‘c’. As
originally described I. americanus has higher value for
the ratio ‘a’ and lower value for the ratio ‘c’ compared
to I. ignavus (a = 34.0–37.0, 40.0–55.0, c = 13.0–15.0,
6.0–10.0, respectively in I. americanus and I. ignavus)
(Bastian, 1865; Cobb, 1914). Our calculation of the
ratio ‘c0 ’ from Cobb’s formula for the species gives a
value of 5.7, while from Bastian’s original drawings it
is about 10.0 for I. ignavus. Although the values for the
ratios ‘c’ and ‘c0 ’ could be useful to differentiate the
two species, the usefulness of the reported slight dif-
ference in the value for the ratio ‘a’, as seen in the light
of the variation of the body width in the genus based
on reproductive maturity, is questionable. For instance,
Tsalolikhin (1987) has reported a range 38.0–63.0 for
the ratio ‘a’ in I. ignavus. Our specimens in the above-
mentioned ratios, in addition to other characteristics,
come closer to I. ignavus, therefore are identified as
such.
This is the first report of the species from Ethiopia.
7. Ironus sphincterus Ebsary, 1985
Measurements
Tables 6 & 7. (Figures 15 & 16)
Description
Population from Zegie
Female:
Body posture after fixation more or less straight (1
specimen) or slightly curved ventrad (3 specimens).
Maximum body width between base of pharynx and
half distance between vulva and anus. Cuticle smooth,
with faint intra-cuticular canals (lateral body pores?)
that were seen as faint lines reaching the outer layer of
the cuticle from inside, mostly conspicuous behind
mid-body; external opening not seen. Metanemes
inconspicuous in most, 2 orthometanemes seen only
in one specimen. Dense crystalloid bodies distributed
in all parts of the body, most concentrated between
base of pharynx and anus, situated probably in the
pseudocoelom as well as in the musculature. Size and
shape of crystalloid bodies variable: 1.0–2.0 m wide
and 2.0–75.0 m long.
An inner circle of six outer labial papillae and an
outer circle of four cephalic setae on the lip region.
Inner labial sensilla inconspicuous, probably papilloid
and small. Outer labial papillae on the anterior side of
lip region. Cephalic setae 27.8–32.1% LRW, situated
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 181

Figure 15. I. sphincterus, female, A: Posterior branch of the reproductive system till the reflexed tip of ovary; B: Part of anterior and complete
posterior branch of reproductive system; C: Pharyngeal region till nerve ring; D: Surface view of anterior end; E: rectal region and tail; F: Entire
female; G: Pharyngo-intestinal junction.

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182

Figure 16. I. sphincterus, male, A: Pharyngeal region till nerve ring; B: Copulatory structures; C: Copulatory structures and tail; D: Entire
male; E: Posterior testis and vas deferens at that level; F: Pharyngo-intestinal junction; G: Spermatozoa.

on the sides of the lip region at the level immediately
anterior to the weak constriction-line between the con-
spicuously bulging lip region and the rest of the body.
Cheilostome short (2.0–3.0 m long) and narrow. Rest
of stoma well sclerotized, very long, and composed
of two parts, an anterior wider chamber followed by a
more or less cylindrical, posterior part. Three almost
equally developed anteriorly curved teeth, one dor-
sal and two ventrosublateral in the wider part of the
stoma. Three pharyngeal gland openings in the stoma,
two anteriorly situated (about 15.0 m from ABE)
ventrosublateral gland openings and one more posteri-
orly (about 54.0 m from ABE) situated dorsal gland
opening. Amphids stirrup shaped, opening situated at

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 183
Table 7. Measurements of males of Ironus sphincterus Ebsary, 1985
refractive sphincter. In female without egg, sphinc-
from Lakes Tana and Ziway.
ter situated 80–110 m from vulva. Vagina muscular;
Ironus sphincterus 44.1–50.0% CBW long. Vulva a transverse slit, situat-
Zegie (L. Tana) L. Ziway ed slightly posterior to mid-body. Eggs 3.3–4.2 times
(n = 4) (n = 1) longer than wide (147–167 m  40.0–45.0 m).
L 2824.8  143.8 (2630.0–3029.0) 3038.0
Tail more or less elongate-conoid, drastically nar-
rowing 65.0–75.0 m from anus; gradually tapering
LRW 15.3  0.5 (15.0–16.0) 16.0
CSL 4.5  0.5 (4.0–5.0) 5.0
more posteriorly, ending in a fine sharp tip; about 5.0
Amph W 9.7  0.5 (9.0–10.0) 8.5 rectal lengths long.
CBW 16.2  0.8 (15.0–17.0) 16.0
Stoma L 101.3  8.3 (95.0–110.0) 108.0 Populations from Gedero and L. Ziway
Stoma W 5.6  0.9 (5.0–6.0) 5.0 One of the two specimens from Gedero was a young
NR 191.3  11.5 (174.0–206.0) 192.0 female with smaller body, narrower amphids, short-
Ph L 549.3  41.2 (498.0–608.0) 536.0 er gonads and vagina, and narrower body. Otherwise
Cardia L 14.3  1.9 (11.0–16.0) 17.0 both specimens were similar to the population from
Cardia W 15.6  2.7 (12.5–20.0) 17.0 Zegie. The population from Ziway was similar to the
CBW 55.5  8.4 (49.0–70.0) 53.0 specimens from Gedero except in having narrower
MBW 56.9  9.9 (50.0–74.0) 54.0 amphids, shorter vagina, narrower body, higher val-
ABW 33.8  1.9 (32.0–37.0) 34.0 ue for the ratio‘a’ and lower value for the ratio ‘Amph
Tail L 189.3  7.1 (182.0–201.0) 200.0 W% CBW’. Specimens from L. Ziway also had more
Spicule L 56.3  6.5 (49.0–65.0) 57.0 intra-cuticular canals (more body pores?).
a 50.9  7.5 (38.6–58.8) 56.3 Male:
b 5.2  0.4 (4.7–5.7) 5.7
c 14.9  0.3 (14.5–15.2) 15.2 Population from Zegie
c0 5.6  0.4 (5.1–6.1) 5.9 Resembling female in many respects except the fol-
NR% 34.9  1.3 (33.7–37.0) 35.8 lowing. Reproductive system diorchic with the poste-
Amph% CBW 59.8  0.7 (58.8–60.6) 53.1 rior testis reflexed posteriad; situated to the right of
CSL% LRW 28.3  3.6 (25.0–33.0) 31.22 the intestine. Spicules typical for the genus, arcuate,
with ventral velum, 1.5–1.8 cloacal body widths long.
Gubernaculum well sclerotized, its distal side sur-
rounding spicule tips. Protractor muscles well devel-
the level of lip region constriction, slightly more than
oped and prominent. Ejaculatory glands not seen. Vas
half corresponding body width wide. Pharynx cylin-
deferens long, extending anterior to mid-body; com-
drical for most of its length with a narrower part at
ponent cells variable in granulation: with fine granules
the level of the nerve ring. Where exactly the pharynx
closer to the cloaca, coarsely granulated more anteri-
musculature starts was not evident, but muscles are
orly. Spermatozoa elongated and narrow, 0.5–1.0 m
unambiguously conspicuous on the dorsal side around
wide and 15.0–20.0 m long, tapering on one side.
the middle of the stoma. Pharyngeal musculature on
Ventral supplement single, setiform, 8.0–10.0 m long,
the ventral side becomes conspicuous about 10.0 m
situated 5.0–6.0 m from cloacal opening.
anterior to the end of the stoma. Nerve ring surrounds
pharynx at about a third of pharyngeal length. Excre-
Specimen from L. Ziway
tory pore seen in one specimen, situated 47.0 m from
Except for the slightly narrower amphids, the speci-
ABE. Ventral gland not seen. Cardia triangular, one-
men from L. Ziway was similar to the population from
third to two-fifths of corresponding body width. Cells
Zegie, L. Tana.
of the intestinal wall filled with brown granules. Intes-
tine leading to 38.0–47.0 m long rectum.
Comparisons and rationale for identification
Female reproductive system didelphic, amphidel-
In having a well sclerotized and refractive sphincter,
phic with reflexed ovaries, position with respect to
our populations come close to three species in the genus
intestine variable. Spermatheca well defined, in some
with a similar character viz. I. elegans Colomba &
fully packed with spermatozoa. Uterus separated from
gonoduct by strongly muscled, well sclerotized and

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184
Vinciguerra, 1979, I. ernsti Argo & Heyns, 1972, and
I. sphincterus Ebsary, 1985.
A. – I. elegans
The original description of I. elegans was based on
one female and two juveniles (Colomba & Vinciguerra,
1979). Their original description as well as our exami-
nation of the holotype shows that our populations differ
from I. elegans in mainly two important aspects i.e. the
tail and amphids. The amphids in I. elegans compared
to their corresponding body width are far narrower than
they are in our populations (Amph W% CBW = 30.0 in
I. elegans). The tail in I. elegans is longer (256.0 m)
and narrows drastically at 20.0% tail L (55.0 m from
anus), posterior part being filiform. Moreover our pop-
ulations have a higher value for the ratio‘c’ (= 9.1 in
I. elegans).
B. I. ernsti
I. ernsti is described as having a bifid dorsal tooth
and inconspicuous nerve ring. Also, females are char-
acterized by having longitudinal vulva and males with
single, reflexed testis. Furthermore, males have wider
lip region and longer ventromedian supplement than
in our populations (LRW = 18.3–19.8 m, supplement
L = 11.0–17.0 m long in males of I. ernsti). These
characteristics make I. ernsti different from our popu-
lations.
C. – I. sphincterus
Our populations are similar to I. sphincterus in
all morphological and morphometrical characteristics
except for the slightly higher value for the ratio ‘c0 ’ in
the Ethiopian populations (c0 = 4.9–5.8 in original pop-
ulation of I. sphincterus) and the absence of males in
the original description. Therefore, we have identified
our specimens as such.
This is the first report of males in the species and
of the species out of its type locality, Canada.
Acknowledgments
We thank Miss R. Van Driessche and Mr M. Bruyneel
for their technical assistance. The first author extends
thanks to The University of Ghent, Belgium, and Bahir
Dar Teachers’ College, Ethiopia, for the study subsidy
and leave, respectively.
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